Taxonomic revision and distribution of the freshwater crabs of the family Pseudothelphusidae (Decapoda, Brachyura) from Guatemala, with the descriptions of two new species and keys to the species of all genera

Directeur De la publication / Publication director : Gilles Bloch Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief : Laure Desutter-Grandcolas assistante De réDaction / assistant editor : Anne Mabille ([email protected]) Mise en page / Page layout : Anne Mabille coMité scientifique / scientific board : Nesrine Akkari (Naturhistorisches Museum, Vienne, Autriche) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Serge Gofas (Universidad de Málaga, Málaga, Espagne) Sylvain Hugel (CNRS, Université de Strasbourg, France) Marco Isaia (Università degli Studi di Torino, Turin, Italie) Rafael Marquez (CSIC, Madrid, Espagne) Jose Christopher E. Mendoza (Lee Kong Chian Natural History Museum, Singapour) Annemarie Ohler (MNHN, Paris, France) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) couverture / cover : Phrygiopilus senticosus n. sp., holotype, UVGCR 31, habitus, dorsal view. Zoosystema est indexé dans / Zoosystema is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Agriculture, Biology, and Environmental Sciences® – Scopus® Zoosystema est distribué en version électronique par / Zoosystema is distributed electronically by: – BioOne® (http://www.bioone.org) Les articles ainsi que les nouveautés nomenclaturales publiés dans Zoosystema sont référencés par / Articles and nomenclatural novelties published in Zoosystema are referenced by: – ZooBank® (http://zoobank.org) Zoosystema est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris / Zoosystema is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Geodiversitas, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol. Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 581 ZOOSYSTEMA • 2025 • 47 (26) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:484420FE-9576-49AF-9049-0CAE5C6AC76E Magalhães C. & Wehrtmann I. S. 2025. — Taxonomic revision and distribution of the freshwater crabs of the family Pseudothelphusidae (Decapoda, Brachyura) from Guatemala, with the descriptions of two new species and keys to the species of all genera. Zoosystema 47 (26): 581-615. https://doi.org/10.5252/zoosystema2025v47a26. http:// zoosystema.com/47/26 ABSTRACT The taxonomic status of several of the freshwater crab species from Guatemala is somewhat confused due to old and poor descriptions of some taxa, description of several species based on female specimens, synonymization and revalidation of species and subspecies, and new combinations made by different authors. Here, we revised the taxonomy and geographic distribution of the Pseudothelphusidae Ortmann, 1893 freshwater crabs from Guatemala, Central America. Historical material deposited in major collections of several institutions was examined, as well as previously unstudied material from the crustacean collection of the Universidad del Valle de Guatemala in Guatemala City. The pseudothelphusid crab fauna of Guatemala consists of 17 currently valid species assigned to six genera: Elsalvadoria Bott, 1967 (two species), Phrygiopilus Smalley, 1970 (five species), Potamocarcinus H.Milne Edwards, 1853 (two species), Raddaus Pretzmann, 1965 (three species), Typhlopseudothelphusa Rioja, 1953 (two species), and Zilchia Pretzmann, 1968 (three species). Five species described only from female specimens are treated as incertae sedis: they are too poorly defined to be classified reasonably. Two species, one from Phygiopilus and the other from Zilchia, are described as new. Illustrations of the first male gonopod and a keys to the species of all genera are provided, as well as a discussion of their geographic and altitudinal distribution accompanied by maps of the species occurrences. Célio MAGALHÃES Laboratório de Bioecologia e Sistemática de Crustáceos (LBSC), Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto (FFCLRP), Universidade de São Paulo (USP), Avenida Bandeirantes, 3900; 14040-901, Ribeirão Preto, SP (Brazil) and Instituto Nacional de Pesquisas da Amazônia, Avenida André Araújo 2936, 69067-375 Manaus, Amazonas (Brazil) [email protected] (corresponding author) Ingo S. WEHRTMANN Museo de Zoología of the Centro de Investigación en Biodiversidad y Ecología Tropical (CIBET), Universidad de Costa Rica, 11501-2060 San José (Costa Rica) and Unidad de Investigación Pesquera y Acuicultura (UNIP) of the Centro de Investigación de Ciencias del Mar y Limnología (CIMAR), Universidad de Costa Rica, 11501-2060 San José (Costa Rica) and Editorial Universitaria, Universidad del Valle de Guatemala, 18 Avenida 11-95, Zonas 15, Vista Hermosa III, Ciudad de Guatemala (Guatemala) [email protected] Submitted on 24 September 2024 | Accepted on 12 March 2025 | Published on 23 October 2025 Taxonomic revision and distribution of the freshwater crabs of the family Pseudothelphusidae (Decapoda, Brachyura) from Guatemala, with the descriptions of two new species and keys to the species of all genera KEY WORDS Biodiversity, Central America, crustacean collections, faunal survey, Neotropical region, zoogeography, new species. 582 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. RÉSUMÉ Révision taxonomique et distribution des crabes d’eau douce de la famille des Pseudothelphusidae (Decapoda, Brachyura) du Guatemala, avec la description de deux nouvelles espèces et une clé des espèces de tous les genres. Le statut taxonomique de plusieurs espèces de crabes d’eau douce du Guatemala est quelque peu confus en raison de descriptions anciennes et médiocres de certains taxons, de plusieurs espèces basées sur des spécimens femelles, de la synonymisation et de la revalidation d’espèces et de sous-espèces, ainsi que de nouvelles combinaisons faites par différents auteurs. Nous avons ici révisé la taxonomie et la répartition géographique des crabes d’eau douce Pseudothelphusidae Ortmann, 1893 du Guatemala, en Amérique centrale. Nous avons examiné des spécimens historiques déposés dans les principales collections de plusieurs institutions, ainsi que du matériel non étudié auparavant provenant de la collection de crustacés de l’Universidad del Valle, à Guatemala City. La faune de crabes pseudothelphusidés du Guatemala comprend 17 espèces actuellement valides, réparties en six genres: Elsalvadoria Bott, 1967 (deux espèces), Phrygiopilus Smalley, 1970 (cinq espèces), Potamocarcinus H. Milne Edwards, 1853 (deux espèces), Raddaus Pretzmann, 1965 (trois espèces), Typhlopseudothelphusa Rioja, 1953 (deux espèces) et Zilchia Pretzmann, 1968 (trois espèces). Cinq espèces décrites uniquement à partir de spécimens femelles sont considérées comme incertae sedis : elles sont trop mal définies pour être classées de manière cohérente. Deux espèces, l’une de Phygiopilus et l’autre de Zilchia, sont décrites comme nouvelles. Des illustrations du premier gonopode mâle et une clé des espèces de tous les genres sont fournies, ainsi qu’une discussion sur leur distribution géographique et altitudinale agrémentée de cartes de répartition des espèces. MOTS CLÉS Amérique centrale, biodiversité, collections de crustacés, inventaire faunique, région néotropicale, zoogéographie, espèces nouvelles. INTRODUCTION Guatemala forms part of the Central American Isthmus, a land bridge that permitted the interchange of North and South American biota and which fostered the evolution of a highly diverse fauna in this region (Bacon etal. 2015; O’Dea etal. 2016). The Central American region harbors a rich fauna of freshwater crabs, which is comprised predominantly by representatives of Pseudothelphusidae Ortmann, 1893 (e.g., Smalley 1970; Pretzmann 1972, 1978, 1980; Rodríguez 1982; Rodríguez& Magalhães 2005; Lara etal. 2013; Magalhães etal. 2013, 2015; Wehrtmann etal. 2016; Magalhães& Ng 2019; Mantelatto etal. 2024) and just a few records of species belonging to Trichodactylidae H.Milne Edwards, 1853 (Rathbun 1893; Pretzmann 1968; Magalhães& Türkay 1996, 2008; Tejeda-Mazariegos etal. 2018; Magalhães& Ng 2019). The primary freshwater crab fauna of Guatemala consists mostly of Pseudothelphusidae species (Cumberlidge etal. 2014). The taxonomic status of several of these species is somewhat confused due to old and poor description of some taxa, descriptions of several species based on female specimens, synonymization and revalidation of species and subspecies, and new combinations made by different authors that have dealt with this group (A.Milne-Edwards 1866; Rathbun 1897, 1905; Pretzmann 1965, 1971, 1972, 1975, 1978, 1980; Bott 1967, 1970; Smalley 1970; Bott 1967, 1970; Delamare Deboutteville 1976a, b; Delamare Deboutteville& Juberthie 1976; Rodríguez 1982; 2001; Rodríguez& Hobbs 1989). Cumberlidge etal. (2014) reviewed the conservation status of the Neotropical freshwater crabs and reported six genera and 12 species for Guatemala. In a more recent update of the diversity and conservation status of the country’s fauna, however, Wehrtmann etal. (2016) listed 17 valid species, including two yet undescribed ones. The present study aimed to revise the taxonomy of the pseudothelphusid crabs from Guatemala, offers high-quality illustrations of their gonopods, describes the two new species mentioned in Wehrtmann etal. (2016), and provides keys for their identification. MATERIAL AND METHODS Taxonomy The material examined herein is deposited in the crustacean collections of the following institutions: American Museum of Natural History, New York; Field Museum of Natural History, Chicago; Instituto Nacional de Pesquisas da Amazônia, Manaus; Muséum national d’Histoire naturelle, Paris; Museo de Zoología, Centro de Investigación en Biodiversidad y Ecología Tropical, Universidad de Costa Rica, San José; National Museum of Natural History, Smithsonian Institution, Washington, D.C.; The Natural History Museum of the United Kingdom, London; Naturhistorisches Museum Wien, Vienna; Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main; Tulane Museum of Natural History, New Orleans; and Universidad del Valle de Guatemala, Guatemala City. Specimens were also obtained from field surveys in Guatemala conducted in January and July 2014. Sampling and transportation were done according to the permit issued by CONAP # 177/2014. For the taxonomic identification we used the keys, descriptions, and illustrations available in Bott (1956, 1970), Smalley (1970), Pretzmann (1972, 1978, 1980), Rodríguez (1982, 2001), and Rodríguez& Hobbs (1989), and by comparison with the type material when available. The description of the gonopod morphology is based on the terminology proposed by Smalley (1964), Rodríguez (1982), and Villalobos& Alvarez (2010). In addition to the nominal names, the entries in the synonymic list cover explicitly or 583 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) implicitly records from Guatemala only. We followed the classificatory system of Álvarez etal. (2020), which also contains keys to the subfamilies. Genera can be identified using the keys in Rogers etal. (2020); keys to the species are offered before the taxonomic treatment of each genus. The label data of some lots have been kept in the original language (Spanish) as much as possible. The meanings are: aldea, village; carretera, road; finca, farm; lago, lake; laguna, lagoon; quebrada, stream; río, river; sierra, mountain range. GeoGraphic disTribuTion The distribution maps were generated using the QGIS software, v. 3.16 (http://www.qgis.org/ptBR/site/); shape files for the hydrographic regions were obtained from the HidroBASINS website (https://www.hydrosheds.org/products/ hydrobasins). The geographic coordinates and elevation data for each record were obtained either from a GPS apparatus during sampling or from data on the labels of the collection lots. When coordinates were unavailable, the localities were georeferenced using Google Earth® software, online gazetteers (GeoNames: http://www.geonames.org/), and selected charts of the Guatemala Topographic Maps 1:50,000 prepared by the Instituto Geográfico Nacional “Ingeniero Alfredo Obiols Gómez”, Guatemala, and the U.S. National Imagery and Mapping Agency, Series E754 (available at https://maps.lib. utexas.edu/maps/topo/guatemala/, accessed in June 2024). Elevation data not available on labels were estimated by plotting geographic coordinates of the place of occurrence in Google Earth® and reading the corresponding elevation at the specific point. In this case, these data are indicated in square brackets. measuremenTs Measurements of the crab carapace are in millimeters when they are available; carapace width was measured across the carapace at its widest point, and carapace length was measured along the midline, from the frontal to the posterior margin. Both measurements follow the number and sex of the specimens examined and are given between parenthesis according to the following format: (carapace width:carapace length); in lots with more than three specimens, only the measurements of the smallest and the largest individuals are informed. abbreviaTions Institutions AMNH American Museum of Natural History, New York; FMNH Field Museum of Natural History, Chicago; INPA Instituto Nacional de Pesquisas da Amazônia, Manaus; NHMB Naturhistorisches Museum Basel, Basel; MNHN Muséum national d’Histoire naturelle, Paris; NHM The Natural History Museum of the United Kingdom, London; NHMW Naturhistorisches Museum Wien, Vienna; SMF Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main; TMNH Tulane Museum of Natural History, New Orleans; UCR-MZ Museo de Zoología, Centro de Investigación en Biodiversidad y Ecología Tropical, Universidad de Costa Rica, San José; USNM National Museum of Natural History, Smithsonian Institution, Washington, D.C.; UVGCR Universidad del Valle de Guatemala, Guatemala City. Other abbreviations used in the text cw carapace width; cl carapace length; elev. elevation; juv juvenile; immat immature; G1, G2 male first and second gonopods, respectively; ov ovigerous; P pereiopod; spec specimen(s); uncat uncatalogued lot. SYSTEMATICS Family pseudoThelphusidae Ortmann, 1893 Subfamily poTamocarcininae Ortmann, 1897 Genus Potamocarcinus H. Milne Edwards, 1853 Potamocarcinus armatus H. Milne Edwards, 1853 (Figs1; 13) Potamocarcinus armatus H. Milne Edwards, 1853: 208; 1854: 174, pl. 13, fig. 3. — Rathbun 1905: 308 (part).— Villalobos Hiriart& Álvarez 2008: 297 (part, in list).— Ng etal. 2008: 176 (part, in list).— Wehrtmann etal. 2016: 779 (in list).— Acevedo-Alonso& Cumberlidge 2022: 582 (Appendix 2, in list). Potamocarcinus (Potamocarcinus) armatus – Bott 1967: 374.— Pretzmann 1971: 20 (in list).— Rodríguez 1982: 119, figs69g, 70a, h, 71c. Potamocarcinus (Potamocarcinus) nicaraguensis aequispinosus Pretzmann, 1971: 20; 1972: 69, figs385, 386, 408, 409. Potamocarcinus (Potamocarcinus) armatus aequispinosus – Pretzmann 1975: 617, pl.2, pl.6 figs18, 19; 1980: 655, pl.15 fig.64. Key To GuaTemalan species of Potamocarcinus h. milne edwards, 1853 1. Carapace with anterolateral margin bearing large, acute teeth (Fig.1A, C); G1 with marginal process and mesodistal lobe of apex erected, not folded over apex cavity (Fig.1G-I); cephalic process vestigial, reduced to two low protuberances (Fig.1I) .................................................................. Po.armatus H. Milne Edwards, 1853 1’. Carapace with anterolateral margin bearing minute, blunt teeth (Fig.2A); G1 with marginal process and mesodistal lobe of apex folded over apex cavity (Fig.2E, F); cephalic process bifid, with two distinct spiniform teeth (Fig.2F) ............................................................................................................ Po.magnus (Rathbun, 1896) 584 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. T ype maTerial examined .— Holotype (by monotypy). Guatemala • 1 ♀ (60.1:40.5), dry; collecting data unknown; MNHNIU-2000-4025 (=MNHN-B4025). oTher maTerial examined.— Guatemala • 1 ♂ 4 ♀; Petén, Laguna de Petén, outlet; [16°58’40.75”N, 89°42’04.80”W; 113m elev.]; 20.II.1935; H. van der Schalie leg.; USNM 189301 • 1juv. ♂ (17.3:11.8) 4 ♀ (20.3:14.4-32.6:22.0); Petén, Lago Petén-Itzá, northern portion; 16°59’48”N, 89°42’10”W; 122m. elev.; 14.I.2014; I. S. Wehrtmann& M. Orozco leg.; UCR-MZ 3234 • 2 ♂, 2 ♀; Petén, Río Subín, Subín; 16°38’14”N, 90°10’59”W; 115m. elev.; 15.I.2014; I. S. Wehrtmann& M. Orozco leg.; UCR-MZ 3235 • AB CD E F G me ms ms Is cst ss me me ce fas mp mp mp cp cp cp HI Fig. 1. — Potamocarcinus armatus H. Milne Edwards, 1853, A, B, female, holotype, dry specimen, MNHN-IU-2000-4025: A, habitus, dorsal view; B, habitus, ventral view; C-I, male, INPA 2123: C, habitus, dorsal view; D, ventral view; E, frontal view; F, left chelae (largest), lateral view; G, G1, left, mesocaudal view; H, left G1, distal portion, cephalic view; I, left G1, distal portion, laterocephalic view. Abbreviations: ce, cephalic process; cp, caudal process; cst, caudal patch of setae; fas, field of apical spines; ls, lateral suture; me, mesial process; mp, marginal process; ms, marginal suture; ss, mesocephalic subdistal patch of setae. Scale bars: A-F, 10 mm; G-I, 1 mm. Photos: C. Magalhães. 585 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) 2 ♂ (33.2:21.7; 41.5:25.9), 1 ♀ (36.3:24.0); Petén, Río Subín, Subín; 16°38’14”N, 90°10’59”W; 115m. elev.; 15.I.2014; I. S. Wehrtmann& M. Orozco leg.; INPA 2123 • 1 ♂ (36.5:23.7) 2 ♀; Alta Verapaz, Río Chajmayic [= Chajmaic] [15°45’05”N, 89°56’45”W; 197m elev.], about midway between Río San Simón and Celba; 14.IV.1935; C.L. Huffs etal. leg.; USNM 230095. disTribuTion.— Guatemala (Alta Verapaz, Petén) (Pretzmann 1980, Wehrtmann etal. 2016; this study) (Fig.13). o Ther records in G uaTemala .— Pretzmann (1980): Petén, Laguna Petén-Itzá, Flores [16°55’39”N, 89°54’10”W; 112m elev.]; Petén, road to Belize, near Macanché [16°57’39”N, 89°38’02”W; 193m elev.]; Petén, Mopan, at Río Melchor Mencos, border with Belize [17°03’27”N, 89°09’06”W; 92m elev.]. descripTion of G1 (Fig. 1G-i) Stout, straight in both caudal, mesial views, moderately compressed mesolaterally; caudal margin nearly straight; lateral margin straight proximally, distinctly concave distally in mesial view; mesial surface with irregular rows of minute setae medially; caudal surface with elongated patch of caudal setae along proximal and median portions, setae longer proximally, decreasing in size medianly; mesocephalic surface with subdistal, elongated patch of setae slightly increasing in size distally. Spermatic channel distal opening on apex cavity, situated towards lateral end at base of caudal process. Marginal suture on mesial surface, straight along most of stem, regularly curved to lateral side distally in mesial view, proximal portion with row of large marginal setae. Marginal process subtriangular, erect, tip slightly rounded, as long as or slightly longer than caudal process in mesial and laterocephalic views, partially overlapping caudal process. Lateral suture incomplete, marked by somewhat deep sulcus on first 3/4 of caudal surface. Mesial process as strong subtriangular projection directed cephalad on distal portion of mesocephalic surface. Apex slightly distorted in cephalic direction; caudal process erect (not folded over apex cavity), largely projected distally, tip rounded in mesial, laterocephalic views. Apex cavity short; field of apical spines weakly developed, with few short, small spines slightly extended distally along half length of cephalic surface of caudal process. Cephalic process vestigial, reduced to two low protuberances subapically on cephalic surface at base of mesial process. remarKs The type locality of this species is unknown (H. Milne Edwards 1853). The holotype of Po.(Po.) nicaraguensis aequispinosus Pretzmann, 1971, a junior synonym of Po.armatus, is a female (NHMB 550a – not examined) from an unknown locality in Guatemala. For the discussion of the taxonomic status and type locality issues regarding these taxa, see Rodríguez (1982: 120). The presumptive occurrence of this species in Mexico was based on specimens from Río Macuspana (Monte Cristo, municipality of Macuspana, state of Tabasco) identified by Rathbun (1905) and Rodríguez& Smalley (1972) as Po. armatus. Pretzmann (1975) used these specimens to describe Potamocarcinus (Po.) hartmanni Pretzmann, 1975. The last species was later recorded from other localities in Tabasco (Villalobos Hiriart& Álvarez 2008). Therefore, it seems reasonable to assume that Po.hartmanni has a northwestern distribution in rivers of the Gulf of Mexico drainage in the Mexican state of Tabasco, whereas Po.armatus is restricted more in the Southeast in rivers of both the Gulf of Mexico and Caribbean drainages in Guatemala and probable Belize as well. Potamocarcinus magnus (Rathbun, 1896) (Figs2; 13) Pseudothelphusa magna Rathbun, 1896: 377, fig.1, 2; pl.29, 30, figs7-10; 1906: 502 (in list). Potamocarcinus guatemalensis Rathbun, 1904: 242 (nomen nudum). Potamocarcinus guatemalensis Rathbun, 1905: 313, text-fig.101, pl.14 fig.7; 1906: 503 (in list).— Coifmann 1939: 110 (in list). Potamocarcinus (Potamocarcinus) magnus guatemalensis – Pretzmann 1965: 2 (in list); 1972: 70, figs381-384, 396-398.— Smalley 1970: 102. Potamocarcinus (Megathelphusa) magnus guatemalensis – Pretzmann 1971: 20 (in list); 1980: 654. Potamocarcinus magnus – Rodríguez 1982: 111, fig.72c, e.— Villalobos F. 1982: 220 (in list).— Ng etal. 2008: 176 (part, in list).— Wehrtmann etal. 2014: 123, fig.1C, D; 2016: 781 (in list).— Acevedo-Alonso& Cumberlidge 2022: 582 (Appendix2, in list). — Villalobos Hiriart& Álvarez 2008: 297 (form, México, Guatemala, El Salvador) (in list). Type maTerial examined.— Holotype. Guatemala • 1 ♂, holotype by monotypy of Potamocarcinus guatemalensis Rathbun, 1905; precise location unknown; M. Bocourt leg.; MNHN Entry 453-1883; MNHN-IU-2017-8050 (= MNHN-B5170). m aTerial examined .— Guatemala • 1 ♂; collecting data unknown; UVGCR 506 • 1 ♂, collecting data unknown; UVGCR 508 • 1 ♂ (65.4:41.6) 1 ♀ (64.9:43.1); Guatemala, Amatitlán lake delta [14°28’34”N, 90°34’35”W; 1190m elev.]; date and leg. unknown; FMNH 1830 • 3 ♂ (43.5:29.0-81.3:-) 1 ♀ (damaged); Guatemala, Amatitlán market; date and leg. unknown; FMNH 1825 • 1 ♂ (82.0:52.0); Guatemala, Lake Amatitlán [14°29’12”N, 90°36’38”W; 1190m elev.]; 28.I-3.II.1906; S. E. Meek leg.; FMNH 1827, • 1 ♂ (74.5:47.9); Guatemala, Lake Amatitlán [14°29’12”N, 90°36’38”W; 1190m elev.]; 28.I-03.II.1906; S.E. Meek leg.; FMNH 1831 • 1 ♂ (56.8:37.3); Guatemala, Lake Amatitlán; 26.VI.1906; S. E. Meek leg.; FMNH 1828 • 3 ♂ (45.3:30.0; 47.4:32.0; 1damaged) 1 ♀ (45.0:30.4); Guatemala, Amatitlán [14°29’12”N, 90°36’40”W; 1190m elev.], date and leg. unknown; FMNH 1832 • 1 immat. ♂; El Progreso, Río Morazán, near Morazán [14°55’54”N, 90°08’21”W; 349m elev.], WNW of El Rancho, Motagua River basin; 8.V.1947; Miller leg.; USNM 1089243 • 1juv. ♂; Izabal, Río Ciénaga [15°40’50”N, 88°57’58”W; 13m elev.], below bridge from road to Sarstún; 18.III.1971; R. M. Boiley, R.C.& J. Lopez leg.; AMNH 8751 • 1juv. ♂; Izabal, Río Ciénaga [15°40’50”N, 88°57’58”W; 13m elev.], below bridge from road to Sarstún; 18.III.1971; R. M. Boiley, R. C.& J. Lopez leg.; AMNH 16670 • 2 ♂ (51.7:37.0; damaged) 1 ♀ (24.9:17.2); Zacapa, Río Palmilla, near Usumatlán; 14°58’30”N, 89°45’45”W; [245m elev.]; 21.VII.2014; I. S. Wehrtmann& M. Orozco leg.; UCR-MZ uncat. • 1 ♂ (28.0:19.5) 2 immat. ♀ (31.7:21.7; 41.3:27.9); Zacapa, Río El Jutillo, near Usumatlán; 14°59’19”N, 89°49’25”W; [347m elev.]; 21.VII.2014, I. S. Wehrtmann& M.Orozco leg.; UCR-MZ uncat. • 1 ♂; Suchitepéquez, Río Choyolate [= Coyolate] [14°22’39.31”N, 586 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. 91°08’13.14”W; 255m elev.], Patulul; 21.V.1948; leg. unknown; USNM 189303 • 1 ♂; Solalá, Lago Atitlán [14°42’N, 91°12’W; 1558m elev.]; 9.X.1993; J. Zunyge leg.; UVGCR 96 • 1 ♂; Solalá, Lago Atitlán; 28.XI.1993; leg. unknown; UVGCR 379 • 1 ♂; Solalá, Lago Atitlán; 6.XII.1984; J. Montúfar leg.; UVGCR 65 • 1 ♂; Solalá, Lago Atitlán; 28.X.1993; E. Pacay leg.; UVGCR 417 • 1 ♂ (70.0:45.0); Solalá, Panajachel (bought in the market, captured at Lago Atitlán); VII.2013; I. S. Wehrtmann leg.; UCR-MZ 3114-02 • 1juv. ♂; Escuintla, carretera a Puerto Quetzal [13°56’N, 90°47’W; 8m elev.]; 1.XI.1993; C. Cardona leg.; UVGCR 319. disTribuTion.— Guatemala (El Progreso, Escuintla, Guatemala, Izabal, Solalá, Suchitepéquez, Zacapa) (Fig.13), Mexico, Belize, Honduras, El Salvador, Costa Rica (Pretzmann 1980; Rodríguez 1982; Villalobos Hiriart& Álvarez 2008; Magalhães etal. 2015; Wehrtmann etal. 2016; this study). oTher records in GuaTemala.— Rodríguez (1982): Escuintla, Aguna [? Río Aguná, 14°15’05”N, 91°07’26”W, 97m elev.]. Pretzmann (1980): Solalá, Lake Atitlán; Zacapa, Rio Cayo, 7.3miles W Estanzuela (128.8km E Guatemala City) [15°00’N, 89°34’W, 190m elev.]; Guatemala, 45km E Guatemala City, Río Platanos [14°50’42.25”N, 90°24’12.67”W, 495m elev.]; Izabal, 2km S Mariscos [15°24’54.53”N, 89°04’43.86”W, 8m elev.], Lake Izabal drainage; Izabal, Río Sena near Seno, 7miles NW Livingston [15°51’12”N, 88°50’53”W, 7m elev.]. The mention of the department of Jalapa in Wehrtmann etal. (2016) must be corrected to department of El Progreso since this is the correct location of the record referred to lot USNM 1089243. descripTion of G1 (Fig. 2c-f) Stout, straight in both caudal, mesial views, moderately compressed mesolaterally; caudal margin nearly straight in mesial view, with tuft of minute to short setae medially; lateral margin slightly concave distally in mesial view; caudal surface with sparce patch of caudal setae along median portion; mesocephalic surface with subdistal, elongated patch of setae increasing in size distally. Spermatic channel opening on apex cavity, situated towards lateral end at base of caudal process. Marginal suture on mesial surface, straight along most of stem, gently curved to lateral side distally in mesial A CD EF ms ss me mp me ce ce cp cp fas Is B Fig. 2. — Potamocarcinus magnus (Rathbun, 1896): A-D, male, MNHN-IU-2017-8050 [holotype of Potamocarcinus guatemalensis Rathbun, 1905] (pereiopods 2-5 missing): A, habitus, dorsal view; B, habitus, ventral view; C, left G1, distal portion, mesial view; D, left G1, distal portion, lateralcephalic view; E, F, male, UCR-MZ 3114-02, left G1, distal portion: E, mesocaudal view; F, laterocephalic view. Abbreviations: ce, cephalic process; cp, caudal process; fas, field of apical spines; ls, lateral suture; me, mesial process; mp, marginal process; ms, marginal suture; ss, mesocephalic subdistal patch of setae. Scale bars: A, B, 10 mm; E, F, 2 mm. Photos: C. Magalhães. 587 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) view, with row of large marginal setae proximally. Marginal process broadly rounded, as long as caudal process in mesial view, partially overlapping this lobe on its lateral side. Lateral suture incomplete, marked by somewhat deep sulcus on first 3/4 of caudal surface. Mesial process as strong subtriangular projection directed cephalad on distal portion of mesial border. Apex moderately distorted in cephalic direction; caudal process enlarged, subquadrate in mesocaudal view, bearing slight protuberance on cephalic end, folded over the apex cavity, partially covering field of apical spines. Apex cavity oblong; field of apical spines moderately developed, rather narrow, with short, small spines. Cephalic process (lateral process, sensu Rodríguez& Hobbs 1989) bifid, with two distinct spiniform teeth apically, both pointing cephalad. remarKs The current geographic distribution of Po.magnus ranges from Mexico to Costa Rica, but some authors (Rodríguez 1982; Villalobos Hiriart& Álvarez 2008) observed morphological differences in the populations of this species inhabiting southern Mexico, Guatemala, and El Salvador from those occurring in Costa Rica. However, at least concerning the G1morphology, the specimen from Lake Atitlán (Fig.2E, F) is very much similar to the holotype specimen from Costa Rica illustrated by Magalhães etal. (2015: 331, figs71, 72), except for the cephalic end of the caudal process, which has a distinct protuberance in the Guatemalan specimen whereas it is angled in the holotype. A more detailed study, including molecular analyses, of this group along its distributional area is needed to clarify this issue. The syntypes of Po.magnus are two males deposited at the USNM (USNM 19048 and 19049) from an unknown locality in Costa Rica. Magalhães etal. (2015) illustrated the G1 of the male USNM 19049 and erroneously considered it as the holotype in the caption of the figures 71 and 72. The holotype of Potamocarcinus guatemalensis Rathbun, 1905, a junior synonym of Po.magnus, is a male (MNHN-IU-2017-8050 [= MNHN-B5170] (Fig.2A-D) from an unknown locality in Guatemala. Subfamily raddausinae Álvarez, Ojeda, Souza-Carvalho, Villalobos, Magalhães, Wehrtmann&Mantelatto, 2020 Genus Elsalvadoria Bott, 1967 Elsalvadoria tomhaasi Bott, 1970 (Figs3A-C; 14) Elsalvadoria tomhaasi Bott, 1970: 361, figs5-8.— Pretzmann 1972: 97, figs492-495, 578-580.— Rodríguez 1982: 95, figs59b, c, e, 61a-d.— Villalobos F. 1982: 220 (in list).— Ng etal. 2008: 173 (in list).— Wehrtmann etal. 2016: 779 (in list).— Acevedo-Alonso& Cumberlidge 2022: 578 (Appendix 2, in list). Elsalvadoria tomasi – Villalobos Hiriart& Álvarez 2008: 295 (in list) [error]. Type maTerial examined.— Holotype. Guatemala • 1 ♂ (41.1:25.1); Sacatepéquez, river near Volcán de Aguas [14°27’54”N, 90°44’35”W]; 900m elev.; date and leg. unknown; SMF 4354. o Ther maTerial examined .— Guatemala • 1 ♂ (25.8:16.5); Guatemala, Mixco [14°38’N, 90°36’W; 1637m elev.]; Finca La Esmeralda; 17.VIII.1993; C. Cardona Gonzalez leg.; UVGCR 344. disTribuTion.— Guatemala (Guatemala, Sacatepéquez) (Bott 1970; Wehrtmann etal. 2016; this study) (Fig.14). The only two available records point to a distribution in the Pacific drainage of southern Guatemala. descripTion of G1 (Fig. 3a-c) Stem moderately sinuous, moderatley compressed in caudocephalic direction, distal portion distinctly curved in laterocephalic direction, slightly wider subdistally; caudal margin, in mesial view, with strong median constriction and enlarged subdistal hump-like lobe, followed by irregular row of variable-sized spines; cephalic margin regularly convex medially in mesial view. Marginal suture on mesial surface, sinuous, gently following curvature of stem, with row of long and short setae along its proximal portion. Marginal process simple, nearly straight in mesial view, not overreaching apex. Lateral suture incomplete, marked by sulcus extending slightly beyond level of median constriction of caudal surface. Mesial process absent. Cephalic surface with cephalic process greatly developed as subdistal unilobed or faintly bilobed enlargement, directed basally and partially twisted towards mesial side, with row of minute spines along its proximal margin. Lateral process enlarged, tongue-shaped; distal margin rounded, situated subdistally on laterocephalic surface, somewhat curved towards cephalic side. Distal portion of lateral surface with distinct rounded (in cephalic view) spiny distolateral lobe. Apex open, oblong, elongated mesolaterally, directed cephalad; mesocaudal margin smooth, produced into small angled lobe, slightly more elevated in relation to laterocephalic margin. Field of apical spines moderately developed, elongated mesolaterally, with small spines. remarKs When comparing the G1 of both examined specimens, we noticed a small variation in the morphology of the cephalic process: in the holotype it is clearly unilobed, with its distal margin broadly rounded (Fig.3C), whereas in the male UVGCR 344 it seems to be faintly bilobed, with its distal margin bearing a slight median concavity. Key To The species of Elsalvadoria boTT, 1967 1. G1 with cephalic process distally unilobed or faintly bilobed (Fig.3A-C); lateral process enlarged, distal margin rounded (Fig.3C) ...................................................................................................... E.tomhaasi Bott, 1970 — G1 with cephalic process distinctly bilobed (Fig.3D-H); lateral process as three-dimensionally conical protuberance (Fig.3E-H) ............................................................................................. E.zurstrasseni (Bott, 1956) 594 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. lobules small, almost indistinct; median groove shallow. Surface of carapace between front and postfrontal lobules smooth, distinctly deflected anteroventrally. Front (Fig.5C, D) with upper border smooth, rounded, median notch absent; lower border carinate, slightly sinuous in dorsal and frontal views, slightly more projected anteriorly than upper one. Upper orbital margin (Fig.5D) lined with faint papillae; lower orbital margin crenulate, lined with faint papillae; exorbital angle very low, obtuse. Anterolateral margin of carapace with set of faint, minute, rounded teeth increasing in size from anterior to posterior portion; posterolateral margin smooth, marked by faint suture. Epistome (Fig.5C, D) narrow longitudinally; epistomial tooth low, slightly deflected anteroventraly, with carinate, smooth borders. Suborbital and subhepatic regions of carapace sidewall (Fig.5C, D) smooth; pterygostomial regions (Fig.5C) weakly pilose. Endopod of Mxp3 (Fig.5C) with ischium subquadrate, its outer and inner margins nearly straight; merus with outer margin broadly rounded, inner surface of palp bearing short and long setae; exopod of third maxiliped long, narrow, 0.73 times length of outer margin of ischium. Aperture of efferent branchial channel (Fig.5D) wide, subretangular, upper margin slightly rounded, smooth. Chelipeds (Fig.5A, B) strongly heterochelous, similarly armed, right P1 larger. Larger cheliped (Fig.5E) with merus subtriangular in cross section; dorsal margin rounded, with irregular row of low tubercles, fainter distally; ventromesial margin lined by longitudinal row of conical tubercles increasing in size distally; ventrolateral margin with single row of faint tubercles; distal upper margin arched, smooth; lower margin with subdistal row of faint tubercles. Carpus with inner margin bearing row of faint tubercles, a prominent median spine, and smooth distally; outer surface rounded, smooth. Palm (Fig.5E) swollen (length/width 1.78), smooth on both mesial and lateral surfaces, ventral margin with irregular row of very faint tubercles. Fingers of larger chela (Fig.5A, B, E) slightly gaping, those of smaller chela not gaping, tips not crossing; pollex with three subtriangular, large teeth interspaced with smaller ones. Dactylus of larger chela moderately arched, approximately as long as palm (dactylus/palm 1.054, measured dorsally); dorsal, mesial and lateral surfaces of dactylus and pollex with longitudinal rows of faint papillae. Ambulatory legs (Fig.5A, B) mostly disarticulated or broken; P5 still attached to body, slender, ratios dactylus/propodus and dactylus/merus 1.40 and 0.79, respectively; P5 with dactylus bearing five longitudinal rows of sharp, corneous spines, increasing in size distally. Thoracic sternum approximately as longer as broad (Fig.5B). Thoracic sternites of 3rd maxillipeds and chelipeds completely fused, except for small notches at lateral edges of sternum. Male sternopleonal cavity densely pilose. Episternites 4-6 triangluar posteriorly, episternite 7 posteriorly truncate. Penis somewhat long and slender, hook-shaped, emerging from nearby coxo-sternal condyle articulation of P5 coxa, located proximally on sternite 8. All pleonal somites free (Fig.5B); lateral margins of somites fringed by line of short setae; male pleon with somite VI trapezoidal, longest, slightly shorter than telson (ration length of telson/length somite VI = 0.92), lateral margins slightly concave. Male telson (Fig.5B) subtriangular, as long as wide, approximately as long as pleonal somite VI, lateral margins nearly straight, fringed by line of short seate, tip rounded. G1 (Fig.5F-H) markedly sinuous, distal portion noticeable curved in laterocephalic direction, compressed in caudocephalic direction, wider proximally in mesocaudal view; caudal margin, in mesocaudal view, with clear median discontinuity due to distinct angled lateral lobe bearing minute setae; cephalic margin broadly convex medianly in mesocaudal view; mesial surface with irregular row of short setae longitudinally along proximal half. Marginal suture on mesial surface, gently following the curvature of the stem, distinctly curved towards lateral side distally in mesial view; row of long and short setae along proximal portion of marginal suture. Marginal process wide, broadly rounded in mesocaudal view, not overreaching mesocaudal margin of apex. Lateral suture incomplete, marked by sulcus along first 2/3 of caudal surface, shallower distally. Mesial process as subtriangular conical spine at base of supraapical process, directed cephalad. Cephalic surface distally and subdistally crenulate, cephalic process absent; mesial surface produced into large, suboval supra-apical process. Supra-apical process with its margins, and all over laterocephalic surface, regularly and densely covered with spines (smaller proximally); cephalic surface distinctly folded proximally, slightly convex in caudocephalic direction medianly; mesocaudal surface smooth, distinctly concave. Supra-apical process slightly wider than long (ratio width/length roughly 1.2), approximately 0.2 length of entire stem in mesocaudal view; tip rounded. Apex narrow, slit-like, elongated mesolaterally, directed cephalad, upper and lower margins finely crenulated. Field of apical spines poorly developed, narrow, elongated mesolaterally, with minute spines nearly concealed by basal portion of supra-apical process and distal margin of cephalic surface of G1. G2 slightly shorter than G1, flagellum very slender, regularly tapering, tip flattened. remarKs In terms of the G1morphology, Ph.senticosus n.sp. is most similar to Ph.strengerae, since their gonopods are strongly recurved cephalically and bear a strong median lobe due to a remarkable discontinuity on the median portion of the caudal margin (better seen in mesocaudal view). However, the following characters can readily distinguish both species: (a) the lateral lobe is less pronounced in Ph.senticosus n.sp., being shorter than the marginal process in mesocaudal view (Fig.5F), whereas this lobe is distinctly more pronounced in Ph.strengerae, being nearly as wide as the marginal process in mesocaudal view (Fig.4J); (b) the width of the stem median constriction is wider in Ph. senticosus n.sp. (ratio width at constriction/width at lateral lobe roughly 0.7 – measured in mesocaudal view) (Fig.5F) than in Ph.strengerae (ratio width at constriction /width at lateral lobe roughly 0.4 – measured in mesocaudal view) (Fig.4J); (c)the mesial process is present in Ph.senticosus n.sp. (Fig.5F) as a conical spine whereas it is absent in Ph.strengerae (Fig.4J); (d) the supra-apical process, particularly towards the mesocaudal surface, is more densely thorny in Ph.senticosus n.sp. (Fig.5F- 595 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) H) than in Ph.strengerae (Fig.4J, K). Additionally, Ph.senticosus n.sp. is distributed in southern Guatemala (department of Escuintla), while Ph.strengerae occurs in the department of Alta Verapaz, in central-northern Guatemala. The new species can be easily differentiated from the other Guatemalan species of Phrygiopilus by the strong lateral lobe and by the cephalic surface of the supra-apical process densely thorny. A C Fsap mp a fas me ms II GH DE B Fig. 5 . — Phrygiopilus senticosus n. sp., holotype, UVGCR 31: A, habitus, dorsal view; B, habitus, ventral view; C, pterygostomial view; D, frontal view; E, right chelae (largest), lateral view; F, left G1, mesocaudal view; G, left G1, caudolateral view; H, left G1, cephalic view. Abbreviations: a, apex; fas, field of apical spines; ll, lateral lobe; me, mesial process; mp, marginal process; ms, marginal suture; sap, supra-apical process. Scale bars: A-E, 10 mm; F-H, 1 mm. Photos: C. Magalhães. 596 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. Genus Raddaus Pretzmann, 1965 Raddaus bocourti (A. Milne-Edwards, 1866) (Figs6A-C; 7; 16) Boscia bocourti A. Milne-Edwards, 1866: 203.— Smith 1870: 146 (in list). Potamocarcinus bocourti – Ortmann 1897: 317 (in list). Pseudothelphusa bocourti – Pocock 1889: 10 (in list).— Rathbun 1898: 513, 533, 537 (in list); 1904: 242 (in list); 1905: 294, pl.15 fig.1; 1906: 502 (in list).— Young 1900: 220.— Coifmann 1939: 106 (in list). Pseudothelphusa similis Rathbun, 1904: 242 (nomen nudum). Pseudothelphusa similis Rathbun, 1905: 295, text-fig.87, pl.15, fig.4; 1906: 502 (in list).— Coifmann 1939: 109 (in list). Pseudothelphusa cobanensis Rathbun, 1904: 242 (nomen nudum). Pseudothelphusa cobanensis Rathbun, 1905: 296, text-fig.88, pl.15, fig.5; 1906: 502 (in list).— Coifmann 1939: 107 (in list).—Pretzmann 1965: 3 (in list).— Smalley 1964: 28, 29, fig.1A, C, E. Pseudothelphusa grallator Rathbun, 1904: 242 (nomen nudum). Pseudothelphusa grallator Rathbun, 1905: 297, text-fig.89, pl.15, fig.6; 1906: 502 (in list).— Coifmann 1939: 108 (in list).— Pretzmann 1965: 3 (in list). Pseudothelphusa magna – Rathbun 1904: 242 (in list); 1905: 299 (part, ? ♀ from Pansos Palochie, Haute Vera Paz); 1906: 502 (in list). Pseudothelphusa cobanensis cobanensis – Bott 1956: 230 (part, typus). Pseudothelphusa (Anaphyrmus) bocourti – Smalley 1965: 651 (in list). Pseudothelphusa (Anaphyrmus) similis – Smalley 1965: 651 (in list). Pseudothelphusa (Anaphyrmus) cobanensis – Smalley 1965: 651 (in list). Potamocarcinus (Raddaus) similis – Pretzmann 1965: 3 (in list). Potamocarcinus (Raddaus) grallator – Pretzmann 1965: 3 (in list). Potamocarcinus (Raddaus) cobanensis – Pretzmann 1965: 3 (in list).— Smalley 1970: 102, fig.14. Potamocarcinus (Raddaus) boccourti [sic] – Pretzmann 1965: 3 (in list). Potamocarcinus (Raddaus) bocourti bocourti – Pretzmann 1971: 20 (in list); 1972: 76, text-figs16, 17, figs442, 443, 453, 698, 699; 1980: 656. Potamocarcinus (Raddaus) bocourti cobanensis – Pretzmann 1971: 20 (in list). Potamocarcinus (Raddaus) bocourti similis – Pretzmann 1971: 20 (in list); 1972: 76, figs447-449, 453-456; 1980: 656. Potamocarcinus (Raddaus) bocourti grallator – Pretzmann 1971: 20 (in list); 1972: 76, figs444-446, 472-474. Raddaus bocourti – Rodríguez 1982: 98, figs64b, e, j, k, 65a-g; 2001: 442.— Villalobos F. 1982: 220 (in list).— Rodríguez& Magalhães 2005: 360 (in table).— Villalobos Hiriart& Álvarez 2008: 297 (in list).— Ng etal. 2008: 177 (in list).— Wehrtmann etal. 2014: 123, fig.1A, B; 2016: 781 (in list).— Acevedo-Alonso& Cumberlidge 2022: 583 (Appendix 2, in list). Type maTerial examined.— Lectotype (by inference of “type” by Rathbun 1905: 294, 295). Guatemala • 1 ♀ (damaged, 76.5:48.5 according to Rodríguez 1982), dry; Alta Verapaz, environs de Cobán [15°27’55.74”N, 90°22’20.10”W; 1303m elev.]; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN Entry 190-1866; MNHN-IU-2000-4052 (= MNHN-B22814). Paralectotype (by inference of “cotype” by Rathbun 1905: 295). Guatemala • 1 ♀ (cw 74.3mm × cl 48.4mm, according to Rodríguez 1982), dry; Alta Verapaz, environs de Cobán [15°27’55.74”N, 90°22’20.10”W; 1303m elev.]; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN-IU-2000-4626 (= MNHN-B22814). Holotype of Pseudothelphusa cobanensis Rathbun, 1905 (by original designation). Guatemala • 1 ♀ (62.5:40.5), dry, ; Alta Verapaz, environs de Cobán; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN Entry 190-1866; MNHN-IU-2000-4625 (=MNHN-B4625, ex B4053). Paratype of Pseudothelphusa cobanensis Rathbun, 1905. Guatemala • 1 ♂ (carapace damaged), dry, ; Alta Verapaz, environs de Cobán; date unknown; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN-IU-2000-4628 (= MNHN-B4628, ex B4053). Holotype of Pseudothelphusa similis Rathbun, 1905. Guatemala • 1 ♂ (70.2:42.5), dry; Alta Verapaz, rivière de Pansos-Palochie [= Polochic river, municipality of Panzós; 15°23’20.74”N, 89°37’30.23”W; 10m elev.]; XII.1865; M. Bocourt leg.; Expédition du Mexique, 1864-1866; MNHN Entry 190-1866; MNHN-IU-2000-4624 (=MNHN-B4624, ex-B4058). Holotype of Pseudothelphusa grallator Rathbun, 1905 (by original designation. Guatemala • 1 ♀ (61.3:38.9), dry, Totonicapán, montagnes des Altos, environs de Totonecapan [? San Francisco El Alto; 14°55’53.07”N, 91°26’42.10”W; 2351m elev.], surrondings of Totonicapán]; M.Bocourt leg.; Expédition du Mexique, 18641866; MNHN-IU-2000-4627 (=MNHN-B4627, ex B4055). Paratypes of Pseudothelphusa grallator Rathbun, 1905. Guate - mala• 1 ♀, dry, 1 ♂ (remnants), dry; same data as in type; MNHNIU-2000-4055 (= MNHN-B4627). o Ther maTerial examined .— Guatemala • 1 ♂; collecting data unknown; UVGCR 46 • 1 ♂; collecting data unknown; UVGCR 209 • 1 ♂, UVGCR 510, collecting data unknown; 1 ♂; Costa Key To GuaTemalan species of raddaus preTzmann, 1965 1. G1 with cephalic surface of distal portion bearing one subdistal spine (Fig.6B, E); mesial process bilobed, enlarged, as wide as or wider than width of main stem in mesial view (Fig.6A, D) ......................................... 2 — G1 with cephalic surface of distal portion bearing two subdistal spines (Fig.6G, H); mesial process unilobed, slightly narrower than width of main stem in mesial view (Fig.6F) ................ R.tuberculatus (Rathbun, 1897) 2. G1 with mesial process trapezoidal, roughly as wide as main stem in mesial view; mesial process with proximal and distal lobes rounded, distal one narrowest (Fig.6A) ....................... R.bocourti (A. Milne-Edwards, 1866) — G1 with mesial process subretangular, wider than main stem in mesial view; mesial process with proximal and distal lobes as subtriangular tooth, distal one smallest (Fig.6D) ............................... R.mertensi (Bott, 1956) 597 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) Sur; 28.X.1993; K. Ness M. leg.; UVGCR 97 • 1 ♂, Costa Sur; XI.1993; C. Cardona leg.; UVGCR 367 • 1juv. ♂ (24.5:16.6); Petén, Poptún, Finca Ixobel, cerca a riachuelo; 16°18’14”N, 89°25’13”W; [537m elev.]; 11.I.2014; I. S. Wehrtmann& M. Orozco leg.; UCR-MZ 3236-01 • 2juv. ♂ (12.4:8.6; 19.8:13.3); Petén, Dolores, Río Sanicté; 16°36’55”N, 89°32’37”W; 270m elev.; 12.01.2014; I. S. Wehrtmann& M. Orozco leg. UCR-MZ 3233 • 1juv. ♂ (21.8:14.5) 1juv. ♀; Quiché, Sacapulas, quebrada El Tesoro, Pacan, 15°18’11”N, 91°04’49”W; 1954m elev.; 20.I.2014; I. S. Wehrtmann, M. Orozco, C. Magalhães& M. W. Dix leg.; UCR-MZ 3232-01 • 2 ♂ spec., Alta Verapaz, Río Chilax, in the surroundings of San Juan Chamelco; [15°26’15.30”N, 90°20’00.89”W; 1368m elev.]; 1975; Böttger leg.; SMF 8866 • 2 ♂ (31.3:20.4; 48.3:30.4) 2 ♀ (22.7:14.7; 42.6:27.1); Izabal, Puerto Barrios, Río Las Escobas; [15°41’09.92”N, 88°38’40.91”W; 93m elev.]; 18.X.1986; leg. unknown; UVGCR 430 • 1 ♂; Izabal, Río Dulce [15°39’26”N, 88°59’58”W; 4m elev.]; 20.X.1997; F. Bolañoz leg.; UVGCR 173 • 1juv. ♂; Izabal, Puerto Santo Tomás [15°41’37”N, 88°37’32”W; 22m elev.], Playa Ramonastes; 29.V.1994; M. L. Priaz& W. Dio leg.; UVGCR 243 • 1 immat. ♂; Izabal, Cerro San Gil, Río Las Escobas [15°39’56.40”N, 88°46’54.72”W; 1113m elev.]; 21.X.2001; Ecología Avanzada – grupo 2 leg.; UVGCR 454 • 2 ♂ (32.8:21.1; 38.8:24.0) 1 ♀ (58.1:35.2); Baja Verapaz, Salamá [15°06’01.88”N, 90°18’51.60”W; 940m elev.]; 20.IV.1934; E. R. Blake leg.; FMNH 1829 • 1 ♂ (33.5:21.4) 3 ♀ (18.8:12.8-27.7:17.5); Zacapa, Villa Rocosa, Río El Jutillo, near Usumatlán; 15°00’45”N, 89°49’58”W; [717m elev.]; 21.VII.2014; I. S. Wehrtmann& M. Orozco leg.; UCRMZ uncat. • 1 ♂; Chimaltenango, Yepocapa [14°30’N, 90°57’W]; 1400m elev.; 6.VI.1949; leg. unknown; USNM 230092 • 1 ♂ 2 ♀ 1juv., Chimaltenango, Yepocapa; 1460m elev.; XI.1949; leg. unknown; USNM 230093 • 1 ♂; Chimaltenango, Yepocapa, 13-14. VII.1949, H. T. Dalmat leg.; USNM 90364 • 1 ♂ (77.4:48.3) 1 ♀ (80.9:49.6), Chimaltenango, stream in area San Pedro Yepocapa; XII.1948; H. T. Dalmat leg.; USNM 88721, • 1 ♂; Guatemala, Lago de Amatitlán, Tacatón; [14°26’27.35”N, 90°33’33.02”W; 1195m elev.]; 31.X.1993; C. MacVean leg.; UVGCR 135 • 1 ♂; Guatemala, Amatitlán; [14°28’29.74”N, 90°34’38.94”W; 1190m elev.]; 6.IV.1991; J. Ruiz R. leg.; UVGCR 148 • 1 ♂; Guatemala, Ciudad de Guatemala, mercado La Plazita [probably captured in Lago Amatitlán]; 14.XI.2000; F. Peloéz leg.; UVGCR 58 • 1 ♂ (74.8:46.6); Guatemala, Santa Catarina Pinula, Puerta Parada [14°33’21.21”N, 90°27’09.28”W; 1837m elev.], Río Acatán; 2.III.1987; W. Dix leg.; UVGCR 523 • 1 ♂; Guatemala, Santa Catarina Pinula, Puerta Parada; 1978; M. Dix leg.; UVGCR 438 • 1 ♂ (83.2:52.3); Guatemala, carretera to El Salvador, km 16; [14°32’47.82”N, 90°27’06.33”W; 1877m elev.]; date unknown; S. Rodas leg.; UVGCR 499 • 1 ♂ (33.4:21.7) 1 ♀ (45.5:29.1); Solalá, boca del río Panajachel [14°44’04.96”N, 91°09’26.20”W; 1565m elev.]; 13.ii.1970; W. Bussing leg.; UCR-MZ 2671-01 • 1 ♂ (70.7:45.8) 1 ♀ (61.1:42.3); Solalá, Lake Atitlán [14°42’N, 91°12’W; 1558m elev.]; date unknown; G.A. Artamonoff leg.; FMNH 12288 • 1 ♂; Solalá, Lago Atitlán; V.1977; G. Martínez leg.; UVGCR 111 • 1 ♂, Solalá, Panajachel [14°44’29”N, 91°09’48”W]; 6.X.1997; J. Cruz leg.; UVGCR 172 • 1 ♂, Solalá, Panajachel, cerca del muelle; [14°44’29.25”N, 91°09’48.20”W; 1563m elev.]; data unknown; J. Cruz leg.; UVGCR 146 • 1 ♂ (65.0:40.0); Solalá, Panajachel (bought in the market, captured at Lago Atitlán); 21.VII.2013; I. S. Wehrtmann leg.; UCR-MZ 3114-03 • 1 ovig. ♀, Solalá, Panajachel (bought in the market, captured at Lago Atitlán); 21.VII.2013; I. S. Wehrtmann leg.; UCR-MZ 3114-01 • 4 ♂ (53.4:33.9-59.0:36.6) 1 ♀ (61.8:38.8); Solalá, Panajachel (bought in the market, captured at lake Atitlán); I.2014; C. Magalhães leg.; INPA 2020 • 1 ♂, Sacatepéquez, Municipalidad Mixco, Finca La Esmeralda [? Finca La Esmeralda, in San Lucas Sacatepéquez, Department Sacatepéquez: 14°34’53.88”N, 90°37’03.71”W; 1637m elev.]; 17.VIII.1999; C. Cardona leg.; UVGCR 16 • 1 ♂ (23.8:16.1); Suchitepéquez, Patutul, Finca Los Tarrales, stream inside the farm; 14°31’28”N, 91°08’07”W; 875m elev.; 19.I.2014; I. S. Wehrtmann, M. Orozco, C. Magalhães& M. W. Dix leg.; UCR-MZ 3231-02 • 15 spec.; Santa Rosa, laguna de Ayarza; [14°25’07”N, 90°07’33”W; 1412m elev.]; 10-11.V.1947; R. R. Miller leg.; USNM 1089230 • 1 ♂; Santa Rosa, Barbarena; [14°18’34”N, 90°21’40”W; 1225m elev]; 9.XI.2000; J. Castillo leg.; UVGCR 171 • 1 ♂; Santa Rosa, Taxisco; [14°04’02”N, 90°28’04”W; 179m elev.]; V.1994; J. R. Valdéz leg.; UVGCR 112 • 2juv. ♂, Santa Rosa, Taxisco-Monterrico; [13°59’38.63”N, 90°27’52.75”W; 30 elev.]; 20.X.2000; leg. unknown.; UVGCR 456 • 1 ♂; Escuintla, Puerto San José; [13°55’45”N, 90°49’13”W; 5m elev.]; 7.XI.2000; D. Maza leg.; UVGCR 390 • 1 ♂; Escuintla, Puerto San José; 21.IX.1997; A. Turcois leg.; UVGCR 159 • 1 ♂; Escuintla, aeropuerto Las Lisas [13°55’43”N, 90°49’53”W; 12m elev.], municipalidad San José; 7.X.1976; M. W. Dix leg.; UVGCR 435. disTribuTion.— Guatemala (Alta Verapaz, Baja Verapaz, Chimaltenango, Chiquimula, Escuintla, Guatemala, Izabal, Petén, Quiché, Santa Rosa, Solalá, Suchitepéquez, Totonicapán, Zacapa) (Fig.16), Mexico, Belize, Honduras, and El Salvador (Pretzmann 1980; Rodríguez 1982, 2001; Villalobos Hiriart& Álvarez 2008; Wehrtmann etal. 2016; this study). o Ther records in G uaTemala .— Pretzmann (1980): Petén, Mopán, at Río Melchor Mencos, border with Belize [17°03’27”N, 89°09’06”W; 92m elev.]; Petén, near Poptún [16°21’N, 89°26’W; 544m elev]; Alta Verapaz, tributary of Río Colochie [= Polochic], NE Tucura [= Tucurú, 15°17’51.32”N, 90°06’58.78”W; 434m elev.]; Chiquimula, Río Apantes [? Aldea Apantes, 14°32’45.73”N, 89°24’28.64”W; 984m elev., municipality of Concepción Las Minas], Melen, NW Esquipulas; Guatemala, 14km S Guatemala City, N San José Pinula [14°32’46”N, 90°24’41”W; 1748m elev.]. Rodríguez (2001): Solalá, Lago de Atitlán, 4miles E of Panajachel [14°41’50.43”N, 91°07’05.52”W; 1604m elev.]; Guatemala, Río Piscaya [= Pixcaya, Department Chimaltenango, 14°51’24.85”N, 90°39’44.99”W; 759m elev.], 2km N junction National RT5-RT4, small tributary of Río Piscaya, at 40km N from Guatemala; Guatemala, km 56 from Guatemala City on route 5 to Salamá, small tributary of Río Grande [El Quiché, 14°53’48.53”N, 90°40’20.42”W; 733m elev.]; Guatemala, km 71 from Guatemala City on route 5 to Salamá, small tributary of Río Grande [Baja Verapaz, 14°54’48.73”N, 90°31’24.78”W; 967m elev.]. descripTion of G1 (Fig. 6a-c) Straight in mesial view, narrow in the proximal 2/3, wide in the distal third, moderately compressed mesolaterally; caudal margin moderately convex subdistally in mesial view; cephalic margin somewhat irregular, with few short setae more widely spaced medianly, close together subdistally at base of widened mesial process. Marginal suture on mesial surface, straight up to apex, with row of large marginal setae along proximal portion partially extending to marginal plate as row of short setae. Marginal process somewhat narrow, slightly rounded, extended as long as mesial margin of apex. Lateral suture incomplete, marked by shallow sulcus on caudal surface. Mesial process widened, nearly as wide as main stem, roughly trapezoidal, directed cephalad, bilobed, with proximal lobe larger, broadly rounded; distal lobe narrow, rounded, situated obliquely in relation to proximal lobe, slightly longer in mesial view; cephalic surface of mesial process distinctly concave in lateral view, convex in mesial view. Apex oblong, situated apically, open, mesial and lateral margins of same height; field of apical spines well developed, with short, small spines; spermatic channel opening on apex cavity, situated towards caudal end. Cephalic surface with strong triangular, subdistal cephalic spine, pointing cephalad. 598 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. remarKs The correct status of the type material of this species and three other synonymous taxa is somewhat unclear. Alphonse Milne-Edwards (1866) offered a very brief description of apparently a single specimen from a river of Coban (Alta Verapaz) for which he provided measurement (cw 75mm× cl 47mm) but did not inform the sex. However, it is not explicitly mentioned in the text that the specimen would be the type (in the sense of the name-bearing specimen). Although it may seem that A. Milne-Edwards (1866) used a single specimen, this cannot be unequivocally assumed because Rathbun (1905) mentioned a slightly smaller female with the same characters in the MNHN collection that she treated as “cotype”. Furthermore, when visiting the MNHN in September 2015 for examining the Guatemalan material, the senior author found a box containing two dry females of R.bocourti (MNHN entry 190-1866 in Bocourt’s “Expédition du Mexique” [catalogued as MNHN-B22814]) with indications of “type” and “co-type”, one of which partially disarticulated and with the carapace badly damaged (Fig.7A). Rathbun (1905) mentioned that “Le type du P. Bocourti, c’est-à-dire l’individu que M. A. Milne-Edwards a mesuré, est une grande femelle” [The type of P. Bocourti, that is to say the individual that Mr. A. Milne-Edwards measured, is a large female] and that “Le femelle cotype est un peu plus petite que le type” [The female cotype is slightly smaller than the type]. By this action, Rathbun (1905) was the first author before the year 2000 to have “unambiguously selected a particular syntype to act as the unique name-bearing type of the taxon” as stated in the Article 74.5 (ICZN 1999), which can be taken as a valid lectotype designation. Later, Rodríguez (1982) measured the larger female with cw 76.5mm × cl 48.5mm, the smaller female with cw 74.3mm × cl 48.4mm, and remarked that the carapace of the former was broken; he treated them as “type” and “paratype”, respectively. Thus, the larger, partially damaged female of MNHN-B22814 A me mp fas cs cs cs me me me DEFG H BC Fig. 6. — Raddaus Pretzmann, 1965 species, G1: A-C, Raddaus bocourti (A. Milne-Edwards, 1866), INPA 2020, left G1: A, mesocaudal view; B, lateral view; C, apical-cephalic view; D, E, Raddaus mertensi (Bott, 1956), holotype, SMF 1822, right G1: D, mesocaudal view; E, laterocephalic view; F-H, Raddaus tuberculatus (Rathbun, 1897), USNM 1089233, left G1: F, mesocaudal view; G, lateral view; H, apical-cephalic view. Abbreviations: cs, subdistal cephalic spine; fas, field of apical spines; me, mesial process; mp, marginal process. Scale bars: 1 mm. 599 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) (Fig.7A) must be considered a valid lectotype designation by Rathbun (1905) and, assuming that A. Milne-Edwards (1866) may have had another specimen at his disposal, the smaller female treated as “cotype” by Rathbun (1905) and kept under the same catalogue number should be treated as paralectotype. A C F EG D B Fig. 7. — Type specimens of Raddaus bocourti (A. Milne-Edwards, 1866) and synonymus species: A, Raddaus bocourti, MNHN-B22814, habitus, dorsal view, female, dry (upper side) and female (damaged), holotype, dry (lower side); B, Pseudothelphusa similis Rathbun, 1905 [synonym of Raddaus bocourti], holotype, male, dry, MNHN-B4624:habitus, dorsal view; C, D, Pseudothelphusa cobanensis Rathbun, 1905 [synonym of Raddaus bocourti], syntypes, habitus, dorsal view: C, female, MNHN-B4625; D, male (damaged), MNHN-B4628; E-G, Pseudothelphusa grallator Rathbun, 1905 [synonym of Raddaus bocourti]: E, female, dry, holotype, MNHN-B4627, habitus, dorsal view; F, male (remnants) (upper side) and female (damaged) (lower side), MNHN-B4627, habitus, dorsal view; G, male (remnants) (lower side) and female (damaged) (upper side), MNHN-B4627, habitus, ventral view. Photos: C. Magalhães. 600 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. Three Pseudothelphusa species from Guatemara were described by Rathbun (1905) based on female specimens: Pseudothelphusa similis, Ps. cobanensis, and Ps. grallator, which have been considered to be junior synomyms of R.bocourti (see Rodríguez 1982; Ng etal. 2008). Rathbun (1905) founded Ps. similis on a single male specimen but also mentioned 17juvenile specimens from the same locality and attributed a slight morphological difference between them to age. As the male, to which she provided measurements, is unequivocally termed “type”, this is the holotype fixed by original designation (ICZN 1999, Art. 73.1.1). The holotype of Ps. similis (MNHN-B4624) is a dry male with a longitudinal cleft along the middle and on the right side of the dorsal region of the carapace (Fig.7B). The type specimens of Ps. cobanensis, a well-preserved female (Fig.7C) and a male in very bad condition (Fig.7D), are also dry. Rathbun’s (1905: 296, 297) action, by clearly treating only the female as “type”, can be interpreted as she has set it as the unique name-bearing type of this taxon (Art. 73.1.1, ICZN 1999), therefore fixing the holotype by original designation. The male was designated as “co-type” (Rathbun, 1905: 297, text-fig.88), which can then be considered as a reference to paratype. This was also Rodríguez’s (1982: 101) interpretation when treated the female as the holotype and the male as paratype. Rathbun (1905: 297) based the description of Pseudothelphusa grallator on at least two specimens: a female (cw 61.3mm × cl 38.9mm) and a male (cw 59.2mm × cl 37.8mm) and, in “Dimensions”, provided a tablewith their measurements. In this table, she indicated the female as “Type”. In the section “Distribution”, she listed eight specimens from the same locality: three females (including the “type”, numbered 623d in the Bocourt’s material from the “Expédition du Mexique”), one male (no.624d in Bocourt’s catalogue), and additionally one female and three young specimens. Since the female with the number 623d (MNHN-B4627) is clearly marked as “Type” by Rathbun (1905), this specimen is the holotype fixed by original designation (Art. 73.1.1, ICZN 1999). Rodríguez (1982) remarked that the situation of these specimens in the Paris Museum is somewhat confusing but, even taking into account the poor condition of the material, he concluded that the reasonably well-preserved female measuring cw 60.5mm × cl 39mm should be the one Rathbun considered as the “Type”. During a visit to the MNHN in September 2015, the senior author found that the condition of some of these specimens had worsened but this dry female was still in reasonably good condition and preserved in a box with the catalogue number 4627 and with a label stating “Bocourt (Expédition du Mexique) (no.623d de son catalogue)”; this box also had the indicative sign of holotype (the red circle label) attached to it (Fig.7E). This specimen should be the one considered by Rodríguez (1982) as the holotype. Another box, bearing a handwritten label and stating “Pseudothelphusa grallator Type 4627”, contained one dry female with the dorsal region of the carapace severely damaged and almost entirely disarticulated as well as the remnants of a male specimen with the abdomen partially broken and the sternopleonal cavity glued to the box (this male has the number “624d” written on the anterior portion of the sternum) (Fig.7F, G). Although these lots bear a label with the word “TYPE” written in red, these specimens should be considered as paratypes; Rodríguez (1982) considered the male as being an allotype. Raddaus mertensi (Bott, 1956) (Figs6D, E; 16) Pseudothelphusa cobanensis mertensi Bott, 1956: 231, pl.30, fig.3ac, pl.35, fig.3a, b. Raddaus mertensi – Cumberlidge etal. 2014: 147 (in table).— Wehrtmann etal. 2016: 781 (in list).— Acevedo-Alonso& Cumberlidge 2022: 583 (Appendix 2, in list). T ype maTerial examined .— Holotype. El Salvador • 1 ♂ (37.3:24.4); Santa Ana, Los Planes Farm [northeast of Metapán, Montecristo National Park; 14°24’N, 89°22’W]; 1830m elev.; 30.X.1950; R.Mertens leg.; SMF 1822. oTher maTerial examined.— Guatemala • 1 ♂ (soft carapace) 3juv. ♀; Chiquimula, Río Atulapa, 3miles east of Esquipulas, on road CA 10; [14°33’45”N, 89°18’37”W; 916m elev.]; 5.IV.1971; R.M. Bailey leg.; USNM 1089229. disTribuTion.— Guatemala (Chiquimula) (Fig.16), Honduras (Ocotepeque) and El Salvador (Santa Ana) (Bott 1956; Wehrtmann etal. 2016; Acevedo-Alonso& Cumberlidge 2022; Magalhães& Wehrtmann 2025; this study). descripTion of G1 (Fig. 6d, e) Stout, straight in mesial view, narrow in proximal 2/3, very wide in distal third, moderately compressed mesolaterally; caudal and cephalic margins straight in mesial view. Marginal suture on mesial surface straight up to apex along most of stem, slightly curved in caudal direction in distal quarter, with short row of large marginal setae along proximal portion. Marginal process somewhat narrow, rounded, not extending beyond mesial margin of apex. Lateral suture incomplete, marked by shallow sulcus on caudal surface. Mesial process greatly widened, slightly wider (roughly 1.2 times) than main stem, subretangular, directed cephalad, bilobed, with proximal lobe bearing broadly subtriangular teeth pointing basally; distal lobe as narrow subtriangular teeth slightly shorter than proximal lobe in mesial view; mesial lobe with proximal margin bearing short row of minute setae; distal margin slightly crenulate, cephalic surface slightly concave in lateral view, bearing strong triangular, subdistal spine pointing cephalad, and small, rounded protuberance located just below subdistal spine. Apex oblong, short, situated obliquely, open, mesial slightly higher than lateral one; field of apical spines moderately developed, with short small spines; spermatic channel opening on apex cavity, situated towards caudal end. remarKs This species was known only from its type locality, in the department of Santa Ana, in northwestern El Salvador. Later, Acevedo-Alonso& Cumberlidge (2022) listed the occurrence of this species in Guatemala and Honduras without specify- 601 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) ing voucher material or distributional data. Thus, our report herein represents the first published record of this species for Guatemala. Raddaus tuberculatus (Rathbun, 1897) (Figs6F-H; 8; 16) Pseudothelphusa tuberculata Rathbun, 1897: 61; 1898: 511 (in key), 513, 535, and 537 (in list); 1904: 242 (in list); 1905: 299, pl.13, fig.9; 1906: 502 (in list).— Young 1900: 217— Coifmann 1939: 109 (in list). Pseudothelphusa (Anaphyrmus) tuberculata – Smalley 1965: 651 (in list). Potamocarcinus (Raddaus) tuberculata – Pretzmann 1965: 3 (in list). Pseudothelphusa (Raddaus) tuberculatus – Pretzmann 1971: 20 (in list); 1972: 80, figs461-464, 519-521. Raddaus tuberculatus – Rodríguez 1982: 103, figs64i, 66a-e.— Villalobos F. 1982: 220 (in list).— Villalobos Hiriart& Álvarez 2008: 297 (in list).— Ng etal. 2008: 177 (in list).— Wehrtmann etal. 2016: 781 (in list).— Acevedo-Alonso& Cumberlidge 2022: 583 (Appendix 2, in list). Type maTerial examined. — Lectotype (designated by Smalley 1965). Guatemala • 1 ♂ (50.0:30.9, in bad condition); Suchite - péquez, stream of Saint Augustín, near Atitlán [? Finca San Agustím; 14°28’23.93”N, 91°12’53.26”W, 609m elev.], Pacific drainage; date unknown, M.Bocourt; USNM 20047. Paralectotype. Guatemala • 1 ♀, Suchitepéquez, stream of Saint Augustín, near Atitlán [? Finca San Agustin; 14°28’23.93”N, 91°12’53.26”W, 609m elev.], Pacific drainage; M.Bocourt leg.; MNHN Entry 844-1866; MNHN-IU-2017-8048 (= MNHNB5322). oTher maTerial examined.— Guatemala • 1 ♂ (69.5:41.4); San Marcos, Finca El Porvenir; [14°57’43”N, 91°56’26”W]; 1370m elev.; 22.III.1952; W. Reeder leg.; USNM 1089233 • 2 ♂ (18.9:13.5-35.1:22.5) 3 ♀ (22.6:14.9-35.7:22.9), San Marcos, sítio San Rafael, San Marcos; 14°56’21”N, 91°54’38”W; 991m elev.; 7.VIII.2015, I. S. Wehrtmann& M. Orozco leg. UCR-MZ uncat. • 1 ♂ (23.5:17.7); Suchitepéquez, carretera a Chicacao; 14°27’50”N, 91°09’01”W; [471m elev.]; 19.I.2014, I. S. Wehrtmann, M. Orozco, C. Magalhães& M. W. Dix leg. UCR-MZ uncat. disTribuTion.— Guatemala (San Marcos, Suchitepéquez) (Fig.16) and Mexico (Rathbun 1897, 1905; Pretzmann 1972; this study). descripTion of G1 (Fig. 6f-h) Somewhat slender, slightly arched in cephalic direction in mesial view, moderately compressed mesolaterally; caudal margin with median constriction in mesial view; cephalic margin gently concave in proximal 2/3, with few short setae subdistally; lateral surface with irregular rows of short to large setae in first half and few minute setae subdistally. Marginal plate fused subdistally to caudal surface. Marginal suture on mesial surface straight up to apex, with row of large to short marginal setae along proximal portion. Marginal process subrectangular, nearly as long as mesial margin of apex. Lateral suture incomplete, marked by distinct sulcus on caudal surface. Mesial process broad, unilobed in mesial view, subtriangular, narrower than width of main stem in mesial view, directed cephalad; proximal corner rounded, distal corner with small sharp spine pointing lateralad in lateral and apical-cephalic views; cephalic surface of mesial process distinctly concave in lateral and apical views. Apex oblong, situated apically, open, mesial and lateral margins about same height; field of apical spines moderately developed, with short small spines; spermatic channel opening on apex cavity, situated medianly. Cephalic surface with two strong, triangular, subdistal spines, pointing cephalad. remarKs Rathbun (1897) described the species based on a male and a female at this time deposited in the holdings of the MNHN (Paris); the male specimen was later sent to the USNM collection (Washington, D.C.). When Smalley (1965) introduced the subgenus Pseudothelphusa (Anaphyrmos), he designated Raddaus tuberculatus as the type species because it was the first species he AB C Fig. 8. — Raddaus tuberculatus (Rathbun, 1897), male, lectotype, USNM 20047: A, habitus, dorsal view; B, habitus, ventral view; C, habitus, detail of the damaged posterior portion of the thoracic sternum. Photos: C. Magalhães. 602 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. included in the subgenus to have been described from a male and, to consolidate his decision, he designated the specimen deposited in the USNM (20047) as lectotype. This specimen is in poor condition, completely disarticulated, the posterior ventral portion severely damaged, the posterior portion of the sternopleonal cavity broken, with both gonopods missing (they were not found in a small vial kept within the jar as usual) (Fig.8). Genus Typhlopseudothelphusa Rioja, 1953 Typhlopseudothelphusa juberthiei Delamare Deboutteville, 1976a (Fig.14) Typhlopseudothelphusa juberthiei Delamare Deboutteville, 1976a: 839, figsG-J.; 1976b: 125, figs6, 7, 14, 15.— Delamare Deboutteville& Juberthie 1976: 23.— Reddell 1981: 14, 17 (in list), 111, fig.16 (map), 113, 321 (in list).— Rodríguez 1982: 123.— Villalobos F. 1982: 220 (in list).— Guinot 1988: 21, fig.13, pl.3 figs3, 4; 1990: 894, fig.6C; 1994: 170.— Ng etal. 2008: 177 (in list).— Villalobos Hiriart& Álvarez 2008: 297 (in list).— Cumberlidge etal. 2014: 144 (in table).— Wehrtmann etal. 2016: 782 (in list).— Acevedo-Alonso& Cumberlidge 2022: 584 (Appendix 2, in list). Typhlopseudotelphusa juberthiei – Cottarelli& Argano 1977: 212 [error]. Typhlopseudothelphusa iuberthiei – Hobbs 1994: 101 (in list) [error]. Type maTerial (noT examined).— Syntypes. Guatemala • 1 ♂; MNHN-IU-2014-23193 (= MNHN-B18869) • 2 ♂; MNHNIU-2014-23194 (= MNHN-B18870) • 1 ♀; MNHN-IU-2014-23192 (= MNHN-B19855) • 1 ♀; MNHN-IU-2014-23196 (= MNHNB18873). Type localiTy.— Guatemala, Alta Verapaz, Chamá mountains, municipality of San Pedro Carchá, Chiacam cave [15°32’49.46”N, 90°06’35.82”W; 858m elev.]. disTribuTion.— Guatemala (Alta Verapaz) (Delamare Deboutteville 1976a, 1976b) (Fig.14). descripTion of G1 See Delamare Deboutteville (1976b). remarKs According to Delamare Deboutteville (1976b), the syntypes were originally deposited at the Laboratoire d’Ecologie Generale of the MNHN. However, these specimens could not be located in the crustacean collection during the visit of the senior author to the MNHN in September 2015 but were subsequently incorporated into this collection (see at https://science.mnhn. fr/institution/mnhn/collection/iu/item/list?fulltext=juberthiei) Typhlopseudothelphusa mitchelli Delamare Deboutteville, 1976a (Fig.14) Typhlopseudothelphusa mitchelli Delamare Debouteville, 1976a: 839, figsD-F; 1976b: 125, figs5, 9-13.— Delamare Deboutteville& Juberthie 1976: 23.— Reddell 1981: 14, 17 (in list), 111, fig.16 (map), 113, 321 (in list).— Rodríguez 1982: 123.— Villalobos F. 1982: 220 (in list).— Guinot 1988: 21, fig.14, pl.3 figs5, 6; 1994: 170, fig.2D, pl.1 fig.8, 9. — Hobbs 1994: 101 (in list).— Ng etal. 2008: 177 (in list).— Villalobos Hiriart& Álvarez 2008: 297 (in list).— Wehrtmann etal. 2016: 782 (in list).— AcevedoAlonso& Cumberlidge 2022: 584 (Appendix 2, in list). Typhlopseudotelphusa mitchelli – Cottarelli& Argano 1977: 212 [error]. descripTion of G1.— See Delamare Deboutteville (1976b). T ype maTerial ( noT examined ).— Syntypes. Guatemala • 1 ♂; MNHN-IU-2017-8057 (= MNHN-B18874) • 2 ♀; MNHNIU-2017-8058 (= MNHN-B19856) • 1 ♀; MNHN-IU-2014-23216 (= MNHN-B18868). Type localiTy.— Guatemala, Alta Verapaz, caves C3 and G3 at Pampur mountains [15°27’38.93”N, 90°35’49.33”W; 906m elev.] disTribuTion.— Guatemala (Alta Verapaz) (Delamare Deboutteville 1976a, 1976b) (Fig.14). remarKs These syntypes were originally deposited at the Laboratoire d’Écologie generale of the MNHN (see Delamare Deboutteville 1976b) but could not be located in the crustacean collection during the visit of the senior author to the MNHN in September 2015. However, they were subsequently incorporated into the crustacean collection (see at http://coldb.mnhn.fr/ catalognumber/mnhn/iu/2017-8057 and http://coldb.mnhn. fr/catalognumber/mnhn/iu/2017-8058). Genus Zilchia Pretzmann, 1968 Zilchia falcata Rodríguez& Hobbs, 1989 (Figs9; 14) Zilchia falcata Rodríguez& Hobbs, 1989: 184, figs1, 2, 3G.—Rodríguez 1992: 184 (in list).— Guinot 1994: 171, pl.1 fig.7.— Hobbs 1994: 101 (in list). —Villalobos Hiriart& Álvarez 2008: 298 (in list).— Cumberlidge etal. 2014: 144 (in table).— Wehrtmann etal. 2016: 782 (in list).— Acevedo-Alonso& Cumberlidge 2022: 584 (Appendix 2, in list). Potamocarcinus falcatus – Ng etal. 2008: 176 (in list). Type maTerial examined.— Holotype. Guatemala • 1 ♂; Alta Verapaz, exact locality unknown; 1975; collected by the Dreux Key To GuaTemalan species of tyPhloPsEudothElPhusa rioJa, 1953 (adapTed from Rodríguez 1982) 1. G1 with mesial process bearing three spines; apex with mesocaudal margin smooth (see fig.5G, H, in Delamare Deboutteville 1976b: 121) ........................................................... T.mitchelli Delamare Deboutteville, 1976 — G1 with mesial process bearing two spines; apex with mesocaudal margin crenulate (see fig.6G, I, in Delamare Deboutteville 1976b: 122) .......................................................... T.juberthiei Delamare Deboutteville, 1976 603 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) Expedition; MNHN-IU-2008-11065 (=MNHN-B20118) as in Rodríguez& Hobbs (1989). Paratype. Guatemala • 1 ♂; MNHN-IU-2014-11972 (=MNHNB20352), as in Rodríguez& Hobbs (1989). oTher maTerial examined. — Guatemala • 1 ♂; Alta Verapaz, exact locality unknown; 1975; collected by the Dreux Expedition; MNHN-IU-2014-10311 (=B27918). disTribuTion.— Guatemala (Alta Verapaz) (Rodríguez& Hobbs 1989) (Fig.14). descripTion of G1 (Fig. 9c, d) Stem moderately compressed mesolaterally, falciform in overall appearance, distal portion distinctly arched cephalad in laterocephalic view; lateral margin convex, caudal margin concave in mesocaudal view; some short seta present along both margins. Caudal surface with small protuberance on median portion of caudal margin; mesial surface with irregular row of few long and short setae longitudinally along proximal half. Marginal suture on mesial surface distinctly curved caudally on distal end, with row of long setae along its proximal portion. Marginal process large, lamelliform in mesocaudal view, not overreaching mesocaudal margin of apex. Lateral suture incomplete, marked by sulcus along first 4/5 of mesocaudal surface, shallower distally. Mesial process (“cephalic process” sensu Rodríguez& Hobbs 1989) greatly developed, strong, subtriangular, long (approximately 1.2 longer than width of apex, measured in mesocaudal view), recurved. Cephalic surface strongly concave distally, with cephalic process as short conical spine (“basal spine” or “lateral process” sensu Rodríguez& Hobbs 1989) situated near base of mesial process; smooth otherwise. Apex oblong, elongated caudocephalicaly; its margins asymmetric, mesocaudal marA B me me CD am al mp cp fas Fig. 9 . — Zilchia falcata Rodríguez & Hobbs, 1989, male, holotype, MNHN-IU-2008-11065: A, habitus, dorsal view; B, habitus, ventral view; C, left G1, mesocaudal view; D, left G1, laterocephalic view. Abbreviations: al, mesocaudal margin of apex; am, laterocephalic margin of apex; cp, cephalic process; fas, field of apical spines; me, mesial process; mp, marginal process. Scale bars: A, B, 10 mm; C, D, 1 mm. Photos: C. Magalhães. Key To GuaTemalan species of Zilchia preTzmann, 1968 1. G1 with apex oblong; field of apical spines well developed, open (not concealed by apical margins); cephalic surface distinctly curved towards cephalic side distally, cephalic lobe absent ................................................ 2 — G1 with apex narrow, slit-like; field of apical spines poorly developed, nearly concealed by apical margins; cephalic surface of distal portion nearly straight distally, with well-developed spiny cephalic lobe (Fig.10A, B ... .......................................................................................................................... Z.germani Pretzmann, 1978 2. G1 with mesial process bearing upper margin serrate; cephalic process as conical spine approximately as large as the mesial process; apex with mesocaudal margin entirely bordered by row of teeth (Fig.11F-I) ................ ............................................................................................................................................ Z.coronata n.sp. — G1 with mesial process bearing upper margin smooth; cephalic process as short conical spine distinctly smaller than mesial process; apex with mesocaudal margin unarmed (Fig.10C, D) ..................................................... .............................................................................................................. Z.falcata Rodríguez& Hobbs, 1989 610 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. Guatemala described solely on female specimens remain uncertain. Smalley (1965) did not include Pseudothelphusa gracilipes A. Milne-Edwards, 1866, Ps. propinqua Rathbun, 1905, and Ps. proxima Rathbun, 1905 in his newly erected subgenus Pseudothelphusa (Anaphyrmos) because of this uncertainty. In addition to these, Rodríguez (1982) also included Ps. bisuturalis Rathbun, 1897 and Gordonia longipes Pretzmann, 1965 among the species he listed as incertae sedis. He argued that, despite being valid names and perhaps even valid species, the impossibility of determining their generic status made their respective taxonomic classification uncertain. Unfortunately, with the material available for this study, we were unable to improve the situation described by Smalley (1965) and Rodríguez (1982). A better understanding of the taxonomic status of these species will only be achieved when a series of additional specimens including adult males from their regions of occurrence to allow a more comprehensive morphological and molecular study. zooGeoGraphy Wehrtmann etal. (2016) counted 11 pseudothelphusid species endemic to Guatemala and this number is kept here. Although Ph.acanthophallus was recently found also in Mexico (Magalhães& Wehrtmann 2025), the current records of Po.armatus show that it is restricted to hydrographic basins of the Caribbean Sea versant of Guatemala. Phrygiopilus acanthophallus does not occur in Nicaragua (see Cumberlidge etal. 2014), but it should occur in Belize as it was found in the Guatemalan side of the Mopan River, which flows in both countries. Guatemala is crossed from west to east by two mountainous ranges that divide the country into the Pacific and the Atlantic hydrographic regions, with the latter further subdivided into river basins flowing north-northwest to the Gulf of Mexico and northeast to the Caribbean Sea (USAID 2002/2003; Suarez 2011). An almost equivalent number of species occur in each of these three hydrographic regions (Fig.18): nine species have been recorded in the Pacific region, eight in the Gulf of Mexico region, and seven in the Caribbean region (Fig.18; Table1). Six species (E.tomhaasi, E.zurstrasseni, Ph. senticosus n.sp., R.mertensi, R.tuberculatus, and Z. germani) are found exclusively in Pacific river basins, whereas the other two regions have three exclusive species each: Ph. chuacusensis, T.mitchelli, and Z.falcata in the Gulf of Mexico basins, and Ph.strengerae, T.juberthei, and Z.coronata n.sp. in the Caribbean basins. Pseudothelphusids are characteristically a montane fauna (Rodríguez 1981, 1982) and the higher diversity and endemicity found in the Pacific region reflects this, as most occurrences are recorded in mountainous areas where the headwaters of rivers that drain into the Pacific are located (Figs14-16). Even in the Gulf of Mexico and Caribbean basins, occurrences are mainly concentrated in the highlands of the country. Species with transbasinal distribution are Ph.acanthophallus and Po.armatus, both occurring in rivers of the Caribbean and Gulf of Mexico hydrographic basins; Ph.ibarrai that is found in the Caribbean and Pacific watersheds; and Po.magnus and R.bocourti are distributed MEXICO 93°W 92°91° 90°89° N 88° 18°N 17° 15° 16° 14° 50 km Po. armatus Po. magnus GUATEMALA BELIZE HONDURAS EL SALVADOR Fig. 13 . — Geographic distribution of Potamocarcinus armatus H. Milne Edwards, 1853 (black triangle) and Po. magnus (Rathbun, 1896) (red circle) in Guatemala (some symbols may represent more than one record). MEXICO GUATEMALA BELIZE HONDURAS EL SALVADOR N 92°93°W 91°90° 89° 88° 18°N 17° 15° 16° 14° 50 km E. zurstrasseni E. tomhaasi T. mitchelli T. juberthiei Z. coronata Z. falcata Z. germani Fig. 14 . — Geographic distribution of Elsalvadoria zurstrasseni (Bott, 1956) (black cross), E. tomhaasi Bott, 1970 (black circle), Typhlopseudothelphusa juberthiei Delamare Debouteville, 1976 (white triangle), T. juberthiei Delamare Deboutteville, 1976 (white circle), Zilchia coronata n. sp. (red circle), Z. falcata Rodríguez & Hobbs, 1989 (red cross), and Z. germani Pretzmann, 1978 (red triangle) in Guatemala. MEXICO GUATEMALA BELIZE HONDURAS EL SALVADOR N 92°93°W 91° 90°89° 88° 18°N 17° 15° 16° 18°N 17° 16° 14° 50 km Ph. acanthophallus Ph. chuacusensis Ph. ibarrai Ph. senticosus Ph. strengerae Fig. 15 . — Geographic distribution of Phrygiopilus acanthophallus Smalley, 1970 (red triangle), P. chuacusensis Smalley, 1970 (red circle), P. ibarrai (Pretzmann, 1978) (black triangle), Ph. senticosus n. sp. (red star), and P. strengerae (Pretzmann, 1965) (black cross) in Guatemala (some symbols may represent more than one record). 611 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) across basins of the three versants (Fig.18; Table1). These last two species have very widespread distributions that encompass Mexico, Belize, El Salvador, Honduras and, in the case of Po.magnus, Costa Rica as well. The mountainous physiography of much of the Guatemalan territory certainly favors a high taxonomic diversification of the pseudothelphusid crabs, which are more diverse and widespread throughout the country than the Trichodactylidae. Trichodactylid crabs, on the other hand, are very little known in the country. There is only one record of an unidentified species of the family in Guatemala, from El Boquerón, near El Estor, department of Izabal (Tejeda-Mazariegos etal. 2018). This family is a group distributed in lowland rivers mainly within the Atlantic drainage (Rodríguez 1981; 1992) and its occurrence in the Lake Izabal region, at 9m elevation, is not surprising. The altitudinal distribution of the Guatemalan pseudothelphusids, obtained from recorded data or inferred from cartographic evidence based on georeferenced data, is presented in Table2. The results indicate that the majority of the pseudothelphusid species occur at elevations above 450m. The various records for Po.magnus and R.bocourti confirm their eurytopic condition also regarding their vertical distribution since they range from nearly sea level to 1558 and 2351m elevation, respectively, in both the Pacific and Caribbean Table 1 . — Presence of the pseudothelphusid species of freshwater crabs from Guatemala in the country’s three major hydrographic regions (*, endemic to Guatemala). Species Caribbean Basin Gulf of Mexico Basin Pacific Basin E. zurstrasseni ■ E. tomhaasi* ■ Ph. acanthophallus ■ ■ Ph. chuacusensis* ■ Ph. ibarrai* ■ ■ Ph. senticosus n. sp.* ■ Ph. strengerae* ■ Po. armatus* ■ ■ Po. magnus ■ ■ ■ R. bocourti ■ ■ ■ R. mertensi ■ R. tuberculatus ■ T. juberthiei* ■ T. mitchelli* ■ Z. coronata n. sp.* ■ Z. falcata* ■ Z. germani* ■ Incertae sedis: B. gracilipes ■ I. longipes ■ Ps. proxima ■ Ps. propinqua ■ Ps. bisuturalis ■ GUATEMALA HONDURAS EL SALVADOR N 92°93°W 91° 90°89° 88° 18°N 15° 17° 18°N 16° 14° 50 km MEXICO R. bocourti R. mertensi R. tuberculatus GUATEMALA BELIZE HONDURAS EL SALVADOR N 92°93°W 91° 90°89° 88° 18°N 15° 17° 18°N 16° 14° MEXICO B. gracilipes I. I. longipes Ps. bisuturalis Ps. propinqua Ps. proxima 50 km Fig. 16 . — Geographic distribution of Raddaus bocourti (A.Milne-Edwards, 1866) (black star), R. mertensi (Bott, 1956) (red circle), and R. tuberculatus (Rathbun, 1897) (red triangle) in Guatemala (some symbols may represent more than one record). Fig. 17 . — Presence of the incertae sedis species in Guatemala: Boscia gracilipes A. Milne-Edwards, 1866 (red triangle), Isabelagordonia longipes Pretzmann, 1965 (red circle), Pseudothelphusa bisuturalis Rathbun, 1897 (white star), Pseudothelphusa propinqua Rathbun, 1905 (white circle), and Pseudothelphusa proxima Rathbun, 1905 (red square). Table 2. — Elevation range of the pseudothelphusid species occurring in Guatemala according to data obtained from the labels of the studied material or inferred from Google Earth® based on the elevation of the estimated location of the occurrence (see Material and Methods). Species Elevation range (m asl) Nr of records Elsalvadoria tomhaasi 900-1637 2 Elsalvadoria zurstrasseni 6-1482 4 Phrygiopilus acanthophallus 929-1382 6 Phrygiopilus chuacusensis 828 1 Phrygiopilus ibarrai 1305-1800 4 Phrygiopilus senticosus n. sp. – No data Phrygiopilus strengerae 805 1 Potamocarcinus armatus 92-197 7 Potamocarcinus magnus 7-1558 15 Raddaus bocourti 4-2351 37 Raddaus mertensi 916 1 Raddaus tuberculatus 471-1370 4 Thyphlopseudothelphusa juberthiei 858 1 Thyphlopseudothelphusa mitchelli 906 1 Zilchia coronata n. sp. 2200 1 Zilchia falcata – No data Zilchia germani 1850-1880 3 612 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. basins (Figs13; 16). Elsalvadoria zurstrasseni also exhibts a wide vertical distribution, ranging from 6 up to 1482m elevation, although most records point to an occurrence in mountainous habitats (Fig.14). The only species not found in elevations higher than 300m is Po.armatus, whose distribution seems to be restricted to the northern lowlands of Guatemala in watersheds of both the Gulf of Mexico and Caribbean hydrographic regions. The revision presented herein contributes to a better understanding of the freshwater crab fauna in Guatemala, but a more realistic picture of the taxonomy as well as horizontal and vertical distributions of the country pseudothelphusid crabs will require a more comprehensive inventory of this group. Acknowledgements We thank the Consejo Nacional de Ciencia y Tecnología, Costa Rica (CONICIT; project IQ-0001-11) and the Conselho Nacional de Desenvolvimento Científico e Tecnologico (CNPq; Brazil, projects 490314/2011-2) for their financial support to ISW and CM. Additional support for ISW was received from Universidad de Costa Rica (projects B2718, B3504, B5655, B6773, and C2721). We are especially grateful to Octavio Cajas (Consejo Nacional de Areas Protegidas de Guatemala) for his support with the collection and export permits. We also thank Christian Birkel (Escuela de Geografía, Universidad de Costa Rica) for helping with the maps. We would like to express our gratitude to Margarita Palmieri and Gabriela Palomo (Biology Department, Universidad del Valle de Guatemala (UVG)), who facilitated the study of the UVG collection, and to Stefania Rodríguez and Christine LeBeau (AMNH), Jochen Gerber (FMNH), Laure Corbari, Paula Martin-Lefrève, and Sébastien Soubzmaigne (MNHN), Peter Dworschak and Martin Schwentner (NHMW), Angelika Brandt, Torben Riehl, Andreas Alspach, Bianca Trautwein, the late Michael Türkay (SMF), Hank Bart Jr. and Nelson Rios (TMNH), as well as Rafael Lemaitre and Karen Reed (USNM) for their assistance during the visits of CM to their respective institutions. A special thanks goes to Mónica Orozco who organized and accompanied us during different field trips in Guatemala, which allowed us collecting specimens reported in this contribution. We also acknowledge the constructive comments of Peter Ng, Paula Martin-Lefèvre, two anonymous referees, and the editors, Laure Desutter-Grandcolas and Anne Mabille, that greatly contributed to improve the manuscript. GUATEMALA BELIZE HONDURAS EL SALVADOR MEXICO Caribbean Basin Gulf of Mexico Basin Pacifie Basin B. gracilipes E. zurstrasseni E. tomhaasi I. longipes Ph. acanthophallus Ph. chuacusensis Ph. ibarrai Ph. senticosus Ph. strengerae Po. armatus Po. magnus Ps. proxima Ps. propinqua Ps. bisuturalis R. tuberculatus R. bocourti R. mertensi T. mitchelli T. juberthiei Z. coronata Z. falcata Z. germani 18°N 94°W 93°92° 91°90° 89°88° 87° 18°N 17° 16° 15° 14° N 50 km Fig. 18. — Geographic distribution of the valid and incertae sedis pseudothelphusid species from Guatemala in the country three major hydrographic regions (Carribean, Gulf of Mexico, and Pacific basins). 613 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) REFERENCES acevedo-alonso a. & cumberlidGe n. 2022.— Conservation status of the neotropical mountain freshwater crabs (Pseudothelphusoidea), in DellaSala D. A.& Goldstein M. I. (eds), Imperiled: The Encyclopedia of Conservation. Volume 2. Elsevier, Amsterdam: 564-589. https://doi.org/10.1016/B978-0-12-821139-7.00091-X Álvarez f., oJeda J. c., souza-carvalho e., villalobos J. l., maGalhães c., wehrTmann i. s. & manTelaTTo f. l. 2020.— Revision of the higher taxonomy of Neotropical freshwater crabs of the family Pseudothelphusidae, based on multigene and morphological analyses. Zoological Journal of the Linnean Society 193 [for 2021]: 973-1001. https://doi.org/10.1093/zoolinnean/zlaa162 b acon c. d., s ilvesTro d., J aramillo c., s miTh b. T., c haKra - barTy p. & anTonelli a. 2015.— Biological evidence supports an early and complex emergence of the Isthmus of Panama. Proceedings of the National Academy of Sciences of the United States of America 112: 6110-6115. https://doi.org/10.1073/pnas.1423853112 boTT r. 1956.— Dekapoden (Crustacea) aus El Salvador. 3. Süsswasserkrabben (Pseudothelphusa). Senckenbergiana biologica 37 (3/4): 229-242. https://decapoda.nhm.org/pdfs/26965/26965.pdf boTT r. 1967.— Fluß-Krabben aus dem westlichen Mittelamerika (Crust., Decap.). Senckenbergiana biologica 48 (5/6): 373-380. https://decapoda.nhm.org/pdfs/26959/26959.pdf boTT r. 1970.— Bemerkungen zu einigen Süsswasserkrabben (Crustacea: Decapoda). Senckenbergiana biologica 51 (5/6): 355461. https://decapoda.nhm.org/pdfs/26953/26953.pdf c audell a. n. 1908.— On some earwings (Forficulidae) collected in Guatemala by Messrs. Schwarz and Barber. Proceedings of the United States National Museum 33 (1563): 169-176. https:// www.biodiversitylibrary.org/page/15668335 coifmann i. 1939.— Potamonidi della Guiana Inglese raccolti dal Prof. Nello Beccari. Archivio Zoologico Italiano 27: 93-116, fig.1-4, pl.3. http://digitale.bnc.roma.sbn.it/tecadigitale/giornale/TO00209791/1939/V.27/00000103 coTTarelli v. and arGano r. 1977.— Trichodactylus (Rodriguezia) mensabak n.sp. (Crustacea, Decapoda, Brachyura), granchio cieco delle acque sotterranee del Chiapas (Messico). Quaderno Problemi Attuali di Scienza e di Cultura, Accademia Nazionale dei Lincei 171(3): 207-212. https://decapoda.nhm. org/pdfs/15030/15030.pdf courbon p. & dreux d. 1976.— Candelaria. Étude du réseau hydrospéléologique de Candelaria – Alta Verapaz. Spelunca, special no.1, Supplément au no.3: 12-17. http://www.chroniques-souterraines.fr/dossiers/Speleologie/01_Explorations/1975guatemala.pdf courbon p., dreux d. & hof b. 1976. — Divers travaux en Alta-Verapaz. Spelunca, spécial no.1, Supplément au no.3: 35-38. http://www.chroniques-souterraines.fr/dossiers/Speleologie/01_Explorations/1975guatemala.pdf cumberlidGe n., alvarez f. & villalobos J. l. 2014.— Results of the global conservation assessment of the freshwater crabs (Brachyura, Pseudothelphusidae and Trichodactylidae): the Neotropical region, with an update on diversity. ZooKeys 457: 133-157. https://doi.org/10.3897/zookeys.457.6598 d elamare d ebouTTeville c. 1976a.— Intérêt biologique et écologique des Crabes cavernicoles du Guatemala et du Mexique appartenant au genre Typhlopseudothelphusa Rioja. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences (Paris) (Série D) 283: 837-840. https://gallica.bnf.fr/ark:/12148/ bpt6k55007634/f925.item d elamare d ebouTTeville c. 1976b.— Sur la radiation évolutive des crabes du genre Typhlopseudothelphusa au Guatémala et au mexique avec description d’espèces nouvelles. Annales de Spéléologie 31: 115-129. https://decapoda.nhm.org/pdfs/32104/32104.pdf delamare debouTTeville c. & JuberThie c. 1976.— Recherches biospéléologiques au Guatemala. Spelunca, spécial Guatémala, Supplément 3: 18-18, 23-24. GuinoT d. 1988.— Les crabes cavernicoles du Monde. Mémoires de Biospéoleologie 15: 3-40, fig.1-17, pl.1-4, tabl. 1-2. https:// www.researchgate.net/publication/312970762_Les_crabes_cavernicoles_du_monde GuinoT d. 1990.— Austinograea alayseaesp. nov., Crabe hydrothermal découvert dans le bassin de Lau, Pacifique sud-occidental (Crustacea Decapoda Brachyura). Bulletin du Muséum national d’Histoire naturelle, Paris, 4e sér., 11, section A (4) [for 1989]: 879-903. https://bionames.org/bionames-archive/issn/0181-0626/11/879.pdf GuinoT d. 1994.— Decapoda Brachyura. In JuberThie C.& d ecou V. (ed.), Encyclopaedia Biospeologica. Tome I. Société de Biospéologie, Moulis (C.N.R.S.) et Bucarest (Académie Roumaine): 165-179, fig.1-3, pl.1. hobbs h. h. Jr., hobbs h. h. iii & daniel m. a. 1977.— A review of the troglobitic decapod crustaceans of the Americas. Smithsonian Contributions to Zoology 244: 1-183. https://doi. org/10.5479/si.00810282.244 hobbs h. h. iii 1994.— Biogeography of subterranean decapods in North and Central America and the Caribbean region (Caridea, Astacidea, Brachyura). Hydrobiologia 287: 95-104. https://decapoda.nhm.org/pdfs/25739/25739.pdf holThuis l. b. 1986.— Decapoda. In boTosaneanu L. (ed.) Stygofauna mundi. A faunistic, distributional and ecological synthesis of the world fauna inhabiting subterranean waters (including the marine interstitial). E.J. Brill, DrW. Backhuys, Leiden: 589-615. https://decapoda.nhm.org/pdfs/29692/29692.pdf. inTernaTional commission of zooloGical nomenclaTure. 1999.— International Code of Zoological Nomenclature (4thed.). International Trust for Zoological Nomenclature. London, U.K. [available online at http://www.iczn.org/iczn/ index.jsp; accessed on 4 March 2025] l ara l. r., w ehrTmann i. s., m aGalhães c. & m anTelaTTo f. l. 2013.— Species diversity and distribution of freshwater crabs (Decapoda: Pseudothelphusidae) inhabiting the basin of the río Grande de Térraba, Pacific slope of Costa Rica. Latin American Journal of Aquatic Research 41: 685-695. https://doi.org/103856/ vol41-issue4-fulltext-5 maGalhães c. & nG p. 2019.— Description of a new genus and species of Pseudothelphusidae Ortmann, 1893 (Decapoda: Brachyura) from Panama, and new records and remarks on the zoogeography of Trichodactylus quinquedentatus Rathbun, 1893 (Trichodactylidae) in Central America. Journal of Crustacean Biology 39(1): 62-73. https://doi.org/10.1093/jcbiol/ruy091 m aGalhães c. & T ürKay m. 1996.— Taxonomy of the Neotropical freshwater crab family Trichodactylidae II. The genera Forsteria, Melocarcinus, Sylviocarcinus, and Zilchiopsis (Crustacea: Decapoda: Brachyura). Senckenbergiana biologica 75 (1/2): 97-130. https://decapoda.nhm.org/references/pdfpick. html?id=11713&pdfroot=https://decapoda.nhm.org/pdfs maGalhães c. & TürKay m. 2008.— Taxonomy of the Neotropical freshwater crab family Trichodactylidae, IV. The genera Dilocarcinus and Poppiana (Crustacea: Decapoda: Brachyura). Senckenbergiana biologica 88 (2): 185-215. https://decapoda. nhm.org/references/pdfpick.html?id=31100&pdfroot=https:// decapoda.nhm.org/pdfs maGalhães c. & wehrTmann i. s. 2025.— Diversity of freshwater crab: new distributional data for four species (Decapoda: Pseudothelphusidae) from Mesoand South American countries. Revista de Biología Tropical 73 (S1): e63625. https://doi. org/10.15517/rev.biol.trop..v73iS1.63625 maGalhães c., campos m. r. & TürKay m. 2013.— Freshwater crabs from eastern Panamá: a new species of Potamocarcinus H. Milne Edwards, 1853, and new records of two little-known species (Crustacea: Decapoda: Pseudothelphusidae, Trichodactylidae). Zootaxa 3702 (4): 348-356. https://doi.org/10.11646/ zootaxa.3702.4.2 maGalhães c., wehrTmann i. s., lara l. r. & manTelaTTo f. l. 2015.— Freshwater crabs from Costa Rica, with a taxonomic revision of the genus Ptychophallus Smalley, 1964 (Crustacea: 614 ZOOSYSTEMA • 2025 • 47 (26) Magalhães C.& Wehrtmann I. S. Decapoda: Pseudothelphusidae). Zootaxa 3905: 301-344. https:// doi.org/10.11646/zootaxa.3905.3.1 m anTelaTTo f. l., m aGalhães c. s ouza -c arvalho e. a., panTaleão J. a. f. & wehrTmann i. s. 2024.— Multi-gene phylogeny of the primary freshwater crab genus Ptychophallus Smalley, 1964 (Pseudothelphusidae: Ptychophallinae) from the Neotropical region. Zoology 164: 126169. https://doi.org/10.1016/j. zool.2024.126169 m ilne -e dwards a. 1866.— Description de trois nouvelles espèces du genre Boscia, crustacés brachyures de la tribu des Thelphuseusiens. Annales de la Société entomologique de France (Série 4) 6: 203-205. https://www.biodiversitylibrary.org/ page/32544981 m ilne e dwards h. 1853.— Mémoire sur la famille des Ocypodiens. Annales des Sciences naturelle (Zoologie), 3e série, 20: 163-228, pls6-11. https://www.biodiversitylibrary.org/page/35466171 milne edwards h. 1854. — Notes sur quelques crustacés nouveaux ou peu connus. Archives du Muséum d’Histoire naturelle 7: 49-192. https://www.biodiversitylibrary.org/page/45672159 newTon e. T. 1892.— On some new reptiles from the Elgin Sandstones. Proceedings of the Royal Society of London 52 (18921893): 389-391. https://www.jstor.org/stable/115310 nG p. K. l., GuinoT d. & davie p. J. f. 2008.— Systema Brachyurorum: Part I. An annotated checklist of extant brachyuran crabs of the world. The Raffles Bulletin of Zoology, Supplement 17: 1-286. https://decapoda.nhm.org/pdfs/27562/27562.pdf o’dea a., lessios h. a., coaTes a. G. Et al. 2016.— Formation of the Isthmus of Panama. Science Advances 2: e1600883. https://doi.org/10.1126/sciadv.1600883 orTmann a. e. 1893.— Die Decapoden-Krebse des Strassburger Museums, mit besonderer Berücksichtigung der von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit im Strassburger Museum aufbewahrten Formen. VII. Theil. Abtheilung: Brachyura (Brachyura genuina Boas) II. Unterabtheilung: Cancroidea, 2. Section: Cancrinea, 1. Gruppe: Cyclometopa. Zoologische Jahrbücher. Abtheilung für Systematik, Geographie und Biologie der Thiere 7(3): 411-495, pl.17. https:// decapoda.nhm.org/pdfs/10704/10704.pdf orTmann a. 1897.— Carcinologische Studien. Zoologische Jahrbücher, Abteilung für Systematik, Geographie und Biologie der Tiere 10: 258-372. https://decapoda.nhm.org/references/pdfpick. html?id=28108&pdfroot=https://decapoda.nhm.org/pdfs p ococK r. i. 1889.— Contributions to our knowledge of the Crustacea of Dominica. The Annals and Magazine of Natural History, series 6, 3: 6-22, pl.2. https://www.biodiversitylibrary. org/partpdf/51330 preTzmann G. 1965.— Vorläufiger Bericht über die Familie Pseudothelphusidae. Sitzungsberichte der MathematischNaturwissenschaftlichen Klasse der Österreichischen Akademie der Wissenschaften, Wien 1: 1-11. https://decapoda.nhm.org/ pdfs/27647/27647.pdf preTzmann G. 1967.— Über einige südamerikanische Süsswasserkrabben (Pseudothelphusidae) Vorläufige Mitteilung. Entomologisches Nachrichtenblatt, Wien 14(2): 23-26. https:// decapoda.nhm.org/pdfs/28113/28113.pdf preTzmann G. 1968.— Weitere neue südamerikanische Süßwasserkrabben (vorläufige Mitteilung). Entomologisches Nachrichtenblatt, Wien 15(2): 1-6. https://decapoda.nhm.org/pdfs/28119/28119. pdf preTzmann G. 1971.— Fortschritte in der Klassifizierung der Pseudothelphusidae. Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Klasse, Abteilung 1, 179 (1-4): 15-24. https:// decapoda.nhm.org/pdfs/27495/27495.pdf p reTzmann G. 1972.— Die Pseudothelphusidae (Crustacea Brachyura). Zoologica 120: 1-182. https://decapoda.nhm.org/ references/pdfpick.html?id=30682&pdfroot=https://decapoda. nhm.org/pdfs preTzmann G. 1975.— Die bedornten Arten der Gattung Potamocarcinus Milne Edwards, 1853. Annalen des Naturhistorischen Museums Wien 79: 615-621. https://www.zobodat.at/pdf/ ANNA_79_0615-0621.pdf preTzmann G. 1978.— Neue Potamocarcinini, Poglayen-Neuwall leg. 1975 (vorläufige Mitteilung). Sitzungsberichte der Österreichischen Akademie der Wissenschaften, Mathematisch Naturwissenchaftliche Klasse, Abteilung 1, 187 (2): 51-54. https://decapoda. nhm.org/pdfs/28121/28121.pdf preTzmann G. 1980.— Von Dr. Ivo Poglayen-Neuwall 1975 in Mittelamerika gesammelte Krabben. Annalen des Naturhistorischen Museums Wien 83: 651-666, pls. 1-18. https://www.zobodat.at/ pdf/ANNA_83_0651-0666.pdf raThbun m. J. 1893.— Descriptions of new species of American fresh-water crabs. Proceedings of the United States National Museum 16 (959): 649-661, pls. LXXIII-LXXVII. https://decapoda.nhm. org/pdfs/31220/31220.pdf raThbun m. J. 1896.— Descriptions of two new species of freshwater crabs from Costa Rica. Proceedings of the United States National Museum 18 [for 1895]: 377-379, pls. 19-30. https:// decapoda.nhm.org/pdfs/27684/27684.pdf raThbun m. J. 1897.— Descriptions de nouvelles espèces de crabes d’eau douce appartenant aux collections du Muséum d’Histoire naturelle du Paris. Bulletin du Muséum nationale d’Histoire naturelle (Paris) 3(1): 58-62. https://www.biodiversitylibrary. org/page/42894843 r aThbun m. J. 1898.— A contribution to a knowledge of the fresh-water crabs of America.—The Pseudothelphusinæ. Proceedings of the United States National Museum 21(1158): 507-537. https://decapoda.nhm.org/pdfs/31221/312 raThbun m. J. 1904.— Les crabes d’eau douce (Potamonidae). Nouvelles Archives du Muséum d’Histoire naturelle, Paris, 4e série, 6: 225-312, pls 9-18. https://www.biodiversitylibrary. org/partpdf/49665 raThbun m. J. 1905.— Les crabes d’eau douce. Nouvelles Archives du Muséum d’Histoire naturelle, Paris, 4e série, 7: 159-322. https:// www.biodiversitylibrary.org/page/36512384 raThbun m. J. 1906.— Catalogue des Potamonidés des collections du Muséum d’histoire naturelle de Paris, d’après les révisions et déterminations de Mlle. Mary Rathbun. Seconde et dernière partie. Bulletin du Muséum nationale d’Histoire naturelle, Paris 12(7): 500-505. https://www.biodiversitylibrary.org/page/5021697 reddell J. r. 1981.— A review of the cavernicole fauna of Mexico, Guatemala, and Belize. The Bulletin of the Texas Memorial Museum 27: 1-327. https://repositories.lib.utexas.edu/bitstreams/5acc4aaa0128-46fa-93a1-217b847f8161/download rioJa e. 1953.— Estudios carcinologicos. XXVIII. Descripcion de un nuevo genero de potamonidos cavernicolas y ciegos de la cueva del Tio Ticho, Comitan, Chis. Anales del Instituto de Biología, Universidad Nacional Autónoma de Mexico, 23(1/2) [for 1952]: 217-225. https://decapoda.nhm.org/pdfs/27333/27333.pdf rodríGuez G. 1981.— Decapoda. In hurlberT s. h., rodríGuez G. & sanTos n. d. (eds), Aquatic Biota of Tropical South America, Part 1: Arthropoda. San Diego State University, San Diego: 41-51. https://decapoda.nhm.org/pdfs/31124/31124.pdf rodríGuez G. 1982.— Les crabes d’eau douce d’Amérique. Familie des Pseudothelphusidae. ORSTOM, Paris, 223p. (Faune Tropicale 22) https://horizon.documentation.ird.fr/exl-doc/pleins_textes/ pleins_textes_6/Fau_trop/01636.pdf rodríGuez G. 1992.— The freshwater crabs of America. Family Trichodactylidae and supplement to the family Pseudothelphusidae. ORSTOM, Paris, 189p. (Faune Tropicale 31) https://horizon. documentation.ird.fr/exl-doc/pleins_textes/divers20-07/36706.pdf rodríGuez G. 2001.— New species and records of pseudothelphusid crabs (Crustacea: Brachyura) from Central America in the Museum of Natural History of Tulane University. Proceedings of the Biological Society of Washington 114: 435-443. https://www. biodiversitylibrary.org/page/35516516 rodríGuez G. & hobbs h. h. Jr. 1989.— A new cavernicol- 615 Pseudothelphusidae from Guatemala ZOOSYSTEMA • 2025 • 47 (26) ous crab, Zilchia falcata, from Guatemala, with notes on the genera of the Potamocarcini (Crustacea Decapoda, Pseudothelphusidae). Bulletin du Muséum national d’Histoire naturelle, 4esérie, 11, SectionA (1): 183-192. https://decapoda.nhm.org/ pdfs/27335/27335.pdf rodríGuez G. & maGalhães c. 2005.— Recent advances in the biology of the Neotropical freshwater crab family of Pseudothelphusidae (Crustacea, Decapoda, Brachyura). Revista Brasileira de Zoologia 22: 354-365. https://www.scielo.br/j/rbzool/a/WDGY K3R9Bq8ZtGtp4S99nkr/?format=pdf&lang=en rodríGuez G. & smalley a. e. 1972.— Los cangrejos de agua dulce de México de la familia Pseudothelphusidae (Crustacea, Brachyura). Anales del Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México 40 (1) [for 1969]: 69-112, pls. 1-12. https://decapoda.nhm.org/ pdfs/27336/27336.pdf roGers c. d.; maGalhães c., peralTa m., ribeiro f. b.; bond-bucKup G., price w. w., Guerrero-KommriTz J., manTelaTTo f. l., bueno a., camacho a. i., GonzÁlez e. r., J ara c. G., p edraza m., p edraza -l ara c., l aTorre e. r. & s anTos s. 2020.— Phyllum Arthropoda: Crustacea: Malacostraca. In d amborenena c., r oGers d. c. & T horp J. h. (eds), “Thorp and Covich’s Freshwater Invertebrates. Fourth Edition - Volume V: Keys to Neotropical and Antarctic Fauna”. Academic Press, London: 809-986. https://doi.org/10.1016/ B978-0-12-804225-0.00023-X s malley a. e. 1964.— A terminology for the gonopods of the American river crabs. Systematic Zoology 13: 28-31. https://doi. org/10.2307/2411434 smalley a. e. 1965.— Two new fresh-water crabs from Nicaragua. The Annals and Magazine of Natural History, series 13, 7 [for 1964]: 651-656. https://decapoda.nhm.org/pdfs/27669/27669.pdf s malley a. e. 1970.— A new genus of freshwater crabs from Guatemala, with a key to the Middle American genera (Crustacea, Decapoda, Pseudothelphusidae). American Midland Naturalist 83: 96-106. https://decapoda.nhm.org/pdfs/27340/27340.pdf smiTh s. i. 1870.— Notes on American Crustacea. No. I. Ocypodoidea. Transactions of the Connecticut Academy of Arts and Sciences 2: 113-176, pls. II-V. https://decapoda.nhm.org/ pdfs/12474/12474.pdf suarez b. G. a. 2011.— Cuencas hidrográficas de Guatemala. Ministerio de Ambiente y Recursos Naturales, Guatemala. Ciudad de Guatemala, 49p. T eJeda -m azarieGos J. c., m eJía o rTíz l. m., l ópez -m eJía m., c randall K. a., p érez -l osada m. & f rausTo -m arTínez o. 2018.— Freshwater crustaceans decadpos: An important resource of Guatemala. In r ay S. (ed.), Biological Resources of Water. IntechOpen, eBook (PDF): 169-179. Available at: https://doi.org/10.5772/intechopen.73638. Accessed on 29 April 2018. usaid - u niTed s TaTes a Gency for i nTernaTional d evelopmenT 2002/2003.— Guatemala Biodiversity and Tropical Forest Assessment. 70p. https://pdf.usaid.gov/pdfdocs/ pnadf213.pdf v illalobos f. a. 1982.— Decapoda. In h urlberT s. h. & v illa - lobos-fiGueroa a. (eds), Aquatic Biota of Mexico, Central America and the West Indies. San Diego State University, San Diego: 215-239. villalobos J. l. & alvarez f. 2010.— Phylogenetic analysis of the Mexican freshwater crabs of the tribe Pseudothelphusini (Decapoda: Brachyura: Pseudothelphusidae). Zoological Journal of the Linnean Society 160: 457-481, 12 figs. https://doi. org/10.1111/j.1096-3642.2009.00606.x villalobos hiriarT J. l. & Álvarez f. 2008.— Los cangrejos de la familia Pseudothelphusidae (Decapoda: Brachyura: Eubrachyura) de México, con un apéndice de las especies citadas para América hasta el 2006. In Álvarez f. & rodríGuezalmaraz G. (eds), Crustáceos de México: Estado Actual de su Conocimiento. Universidad Autónoma de Nuevo León, Monterrey, Mexico: 239-300. w ehrTmann i. s., m aGalhães c. & o rozco m. 2014.— Freshwater crabs in Lake Atitlán, Guatemala: not a single-species fishery. Journal of Crustacean Biology 34: 123-125. https://doi. org/10.1163/1937240X-00002200 w ehrTmann i. s., m aGalhães c. & o rozco m. n. 2016.— An update on the primary freshwater crab fauna of Guatemala (Decapoda, Brachyura, Pseudothelphusidae). Journal Crustacean Biology 36(6): 776-784. https://doi.org/10.1163/1937240X-00002478 younG c. G. 1900.— The stalk-eyed Crustacea of British Guiana, West Indies, and Bermuda. John M. Watkins, London, 514p. https://www.biodiversitylibrary.org/page/11111674 Submitted on24September 2024; accepted on 12 March 2025; published on 23 October 2025.